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Full text of "Transactions of the Royal Society of Edinburgh"

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TRANSACTIONS 



OF THE 



KOYAL SOCIETY OF EDINBURGH. 



' <t . C i'l 



TRANSACTIONS 



OF THE 



ROYAL SOCIETY 



OF 



EDINBURGH. 



VOL. L. 
Nl8T0g! 



EDINBURGH : 

PUBLISHED BY ROBERT GRANT & SON, 107 PRINCES STREET, 
AND WILLIAMS & NORGATE, 14 HENRIETTA STREET, COVENT GARDEN, LONDON. 



MDCCCCXVI. 



ro. i. 


Published 


January 7, 1914. N 


o. XV. 


Published 


May 5, 1915. 


II. 


)> 


December 29, 1913. 


XVI. 


!> 


December 9, 1914. 


in. 


)! 


March 30, 1914. 


XVII.' 


>) 


February 26, 1915 


IV. 


55 


March 30, 1914. 


XVIII. 


55 


March 10, 1916. 


v. 


)) 


March 12, 1914. 


XIX. 


>> 


April 13, 1915. 


VI. 


55 


April 1, 1914. 


XX. 


55 


May 11, 1915. 


VII. 


55 


April 15, 1914. 


XXI. 


55 


May 11, 1915. 


VIII. 


55 


April 30, 1914. 


XXII. 


!) 


May 11, 1915. 


IX. 


)> 


June 4, 1914. 


, XXIII. 


55 


May 20, 1915. 


X. 


>! 


June 30, 1914. 


, XXIV. 


1) 


May 20, 1915. 


XI. 


55 


August 18, 1914. 


XXV. 


51 


May 20, 1915. 


XII. 


55 


July 29, 1914. 


, XXVI. 


5 5 


May 22, 1915. 


XIII. 


5' 


October 3, 1914 


. XXVII. 


55 


June 22, 1915. 


XIV. 


>5 


July 2, 1915. 


, XXVIII 


)' 


August 24, 1915. 



CONTENTS. 



PART I. (1913-14.) 

NUMBER PAGK 

I. Sphserostoma ovale (Conostoma ovale et intermedium, Williamson), 
a Lower Carboniferous Ovule from Petty cur, Fifeshire, Scotland. 
By Margaret J. Benson, D.Sc, Professor of Botany in the Uni- 
versity of London, Head of the Department of Botany at the Royal 
Holloway College, Englefield Green, Surrey. Communicated by 
Dr R. Kidston, F.R.S. (Plates I and II), . . . - . 1 

II. Studies on the Pharmacological Action of Tetra-Alhyl- Ammonium 
Compounds. I. The Action of Tetra-Methyl- Ammonium Chloride. 
By Professor C. R. Marshall, . . . . .17 

III. Polychseta of the family Nereidae, collected by the Scottish National 

Antarctic Expedition {1902-190 A). By L. N. G. Ramsay, M.A., 
B.Sc., Carnegie Research Scholar, Christ's College, Cambridge. 
Communicated by Dr J. H. Ashworth. (Plate III), . . 41 

IV. On the Genus Porponia and Related Genera, Scottish National 

Antarctic Expedition. By Professor Oskar Carlgren, Universitetets 
Zoologiska Institution, Lund. Communicated by Dr W. S. Bruce. 
(Plate IV), 49 

V. On the Fossil Flora of the Staffordshire Coal Fields. Part III. — The 
Fossil Flora of the Westphalian Series of the South Staffordshire 
Coal Field. By R Kidston, LL.D., F.R.S. (Plates V-XVI), . 73 

VI. The Anatomy of a New Species of Bathydoris, and the Affinities of 
the Genus : Scottish National Antarctic Expedition. By T. J. 
Evans, M.A. (Oxon.), Lecturer in Zoology in the University of 
Sheffield. Communicated by Dr J. H. Ashworth. (Plates XVII 
and XVIII), . . . . . . . .191 



vi CONTENTS. 

NUMKKK PAGE 

VII. Rupture Stresses in Beams and Crane Hooks. By Angus R. Fulton, 
B.Sc, A.M.Inst.C.E., Engineering Department, University College, 
Dundee. Communicated by Professor A. H. Gibson, D.Sc, 
A.M.Inst.C.E., ....... 211 

VIII. Scottish National Antarctic Expedition : A Description of the 
Systematic Anatomy of a Fatal Sea- Leopard {Stenorhynchus 
leptonyx), with Remarks upon the Microscopical Anatomy of some 
of the Organs. By Harold Axel Haig, M.B., B.S., M.R.C.S., late 
Lecturer in Histology and Embryology, University College, Cardiff; 
M'Robert Research Fellow, University of Aberdeen. Communicated, 
by Professor Arthur Robinson, M.D. (Plates XIX-XXII), . 225 



PART II. (1913-14.) 

IX. Stalk-ei/ed Crustacea Malacostraca of the Scottish National 
Antarctic Expedition. By Rev. Thomas R. R. Stebbing, M.A., 
F.R.S., F.L.S., F.Z.S., Hon. Fellow of Worcester College, Oxford. 
Communicated by Dr J. H. Ashworth. (Plates XXIII-XXXII), . 253 

X. The Aborigines of Tasmania. Part III. The Hair of the Head 
compared with that of other Ulotrichi and with Australians and 
Polynesia.'n s. By Principal Sir William Turner, K.C.B., D.C.L., 
F.R.^., __l~ht of the Royal Prussian Order Pour le Merite, Emeritus 
Professor of Anatony. (With Figures in Text), . . . 309 

XI. The Pinna-Trace in the Ferns. By R. C. Davie, M.A., B.Sc, late 
Robert Donaldson Research Scholar in the University of Glasgow, 
Lecturer in Botany in the University of Edinburgh. Communicated 
by Professor I. Bayley Balfour, F.R.S. (Plates XXXIII-XXXV), 349 

XII. Studies on the Pharmacological Action of Tetra- Alky i- Ammonium 
Compounds. II. The Action, of Tetra- Ethyl- Am,monium Chloride. 
By Professor C. R. Marshall. (With Figures in Text), . . 379 

XIII. Rocks from Gough Island, South Atlantic [collected by the Scottish 
National Antarctic Expedition, 1902-1904) By Robert Campbell, 
M.A., D.Sc, Lecturer in Petrology in the University of Edinburgh. 
Communicated by Professor James Geikik, D.C.L., LL.D., F.R.S. 
(Plate XXXVI), 397 



CONTENTS. Vll 

NUMBER PAGE 

XIV. On a New Species of Sclerocheilus, with a Revision of the Genus. By 
J. H. Ash worth, D.Sc, Lecturer in Invertebrate Zoology in the 
University of Edinburgh. (Plate XXXVII, and Four Text-figures), 405 

XV. Atlantic Sponges collected by the Scottish National Antarctic 
Expedition. By Jane Stephens, B.Sc. Communicated by Dr 
W.S.Bruce. (Plates XXXVIII-XL), .... 423 

XVI. On the Fossil Osmundaeeas. By R. Kidston, LL.D., F.R.S., F.G.S., 
Foreign Mem. Imper. Mineralogical Society of Petrograd, Hon. Sec. 
R.S.E. ; and D. T. Gwynne-Vaughan, M.A., F.Pv.S.K, M.R.I.A., 
Professor of Botany, University College, Reading. (Plates XLI- 
XLIV), ........ 469 

XVII. Studies on the Pharmacological Action of Tetr a- Alkyl- Ammonium 
Compounds. II I. The Action of Methyl - Ethyl - Ammonium 
Chlorides. By Professor C. R. Marshall, . . . .481 



PART III. (1913-14.) 

XVIII. The Histology of Disseminated Sclerosis. By James W. Dawson, 
M.D., Neurological Histologist to the Royal College of Physicians' 
Laboratory ; formerly Carnegie Research Fellow. To which is 
prefaced a Preliminary Communication on the subject made to the 
Pathological Society of Great Britain and Ireland by the late 
Alexander Bruce, M.D., LL.D., and James W. Dawson, M.D. 
Communicated by A. Ninian Bruce, M.D. (Plates XLV- 
LXXVIII), ........ 517 



PART IV. (1913-14.) 

XIX. Temperature Observations in Loch Earn. Part II. By E. M. 

Wedderburn, D.Sc, and A. W. Young, M. A., B.Sc, . . 741 

XX. On TTsemonais laurentii, n. sp., a Representative of a little-known 
Genus of Naididse. By J. Stephenson, D.Sc, Professor of Zoology, 
Government College, Lahore. (Plate LXXIX), . . .769 



viii CONTENTS. 

Nl'MHKl! PA0E 

XXI. On a Rule of Proportion observed in the Setse of certain Naididce. 
By J. Stephenson, M.B., D.Sc, Professor of Zoology, Government 
College, Lahore, . . . . . • .783 

XX II. On the Sexual Phase in certain of the Naididx. I. The Anatomy of 
Sexual Individuals of the Genus Dero ; with Remarks on Hwmonais. 
II. The Genital Organs in the Genus Slavina. By J. Stephenson, 
D.Sc., Professor of Zoology, Government College, Lahore. (Plate 
LXXX), . 789 

XXIII. Geological Observations in South Georgia. By D. Ferguson, Mem. 

Inst. M.E., F.R.G.S. Communicated by Professor J. W. Gregory, 
D.Sc.,F.R.S. (Plates LXXXI-XCI), . . . .797 

XXIV. The Geological Relations and Some Fossils of South Georgia. By 

J. W. Gregory, D.Sc., F.R.S. (Plates XCII-XCIII), . . 817 

XXV. The Petrology of South Georgia. By G. W. Tyrrell, A.R.C.Sc, 
F.G.S., Lecturer in Mineralogy and Petrology, University of 
Glasgow. (Plate XCIV), . . . . . .823 

XXVI. The Anatomy and Affinity of Deparia Moorei, Hook. By John 
M'Lean Thompson, M.A., B.Sc, Glasgow University. Communicated 
by Professor Bower. (Plates XCV-XCVIi), . . .837 

XXVII. Morphology and Mathematics. By D'Arcy Wentworth Thompson, 857 

XXVIII. The Poisoned Arrows of the Abors and Mishmis of North- East India, 
and the Composition and Action of their Poisons. By Sir Thomas 
R. Fraser, M.D., F.R.S., Professor of Materia Medica and Thera- 
peutics in the University of Edinburgh. (Plates XCVIII-C), . 897 



Index, . x£v^ H < Vu sSx • .931 




UrC. 



TRANSACTIONS 



OF THE 



ROYAL SOCIETY OF EDINBURGH. 

VOLUME L. PART I.— SESSION 1913-14. 




CONTENTS. 

PAOH 

I. Sphserostoma ovale (Conostoma ovale et intermedium, Williamson), a Lower Carboniferous 
Ovule from Petty cur, Fifeshire, Scotland. By Maegarbt J. Benson, D.Sc., Professor of 
Botany in the University of London, Head of the Department of Botany at the Royal 
Hollo way College, Englefield Green, Surrey. Communicated, by Dr R. Kidston, F.R.S. 
(Plates I. and II.), ......... 1 

(Issued January 7, 1914.) 

TT. Studies nn the Pharmacological Action nf Tetra-alkul-ammonium Compounds. I. The Action 

&. Marshall, . . .17 

.3.) 

tish National Antarctic Expedition 
ERRATUM. Carnegie Research Scholar, Christ's 

drth. (Plate III.), . .41 

Line 11 from top, for "p. 785" read " p. 791." *•) 



National Antarctic Expedition. By 
Institution, Lund. Communicated 



-) 



49 



V. On the Fossil Flora of the Staffordshire Coal Fields. Part III. — The Fossil Flora of the 
Westphalian Series of the South Staffordshire Coal Field. By R. Kidston, LL.D., F.R.S. 
(Plates V.-XVL), .......... 73 

(Issued March 12, 1914.) 

VI. The Anatomy of a New Species of Bathydoris, and the Affinities of the Genus: Scottish National 
Antarctic Expedition. By T. J. Evans, M.A. (Oxon.), Lecturer in Zoology in the University 
of Sheffield. Communicated by Dr J. H. Ashworth. (Plates XVII. and XVIII. ), . 191 

(Issued April 1, 1914.) 



* 



VII. Rupture Stresses in Beams and Crane Hooks. By Angus R. Fulton. B.Sc, A.M.Inst.C.E., 
Engineering Department, University College, Dundee. Communicated by Professor A. H. 
Gibson, D.Sc, A.M.Inst.C.E., . . . . . . . .211 

(Issued April 15, 1914.) 

VIII. Scottish National Antarctic Expedition : A Description of the Systematic Anatomy of a Foetal 
\A Sea-Leopard (Stenorhynchus leptonyx), with Remarks upon the Microscopical Anatomy of 
some of the Organs. By Harold Axel Haig, M.B., B.S., M.R.C.S., late Lecturer in 
Histology and Embryology, University College, Cardiff; M 'Robert Research Fellow, 
University of Aberdeen. Communicated by Professor Arthur Robinson, M.D. (Plates 
XIX.-XXIL), .225 

(Issued April SO, 1914.) 



EDINBURGH: 

PUBLISHED BY ROBERT GRANT & SON, 107 PRINCES STREET, 
AND WILLIAMS & NORGATE, 14 HENRIETTA STREET. COVENT GARDEN, LONDON. 



MDCCCCXIV. 
Price Twenty-fivi Shillings and Ninepence. 



viii CONTENTS. 

XXI. On a Rule of Proportion observed in the Setse of certain Naididcs. 
By J. Stephenson, M.B., D.Sc, Professor of Zoology, Government 
College, Lahore, ....... 783 

XXII. On the Sexual Phase in certain of the Naididaz. I. The Anatomy of 
Sexual Individuals of the Genus Dero ; with Remarks on Hwmonais. 
II. The Genital Organs in the Genus Slavina. By -J. Stephenson, 
D.Sc., Professor of Zoology, Government College, Lahore. (Plate 
LXXX), . 789 

XXIII. Geological Observations in South Georgia. By D. Ferguson, Mem. 

Inst. M.E., F.R.G.S. Communicated by Professor J. W. Gregory, 
D.Sc.,F.R.S. (Plates LI 

XXIV. The Geological Relations c 

J. W. Gregory, D.Sc., F. 

XXV. The Petrology of South C 
F.G.S., Lecturer in Mi 
Glasgow. (Plate XCIV), 

XXVI. The Anatomy and Affinity v 

M'Lean Thompson, M.A., B.Sc, Glasgow University. Communicated. 

by Professor Bower. (Plates XCV-XCVI I), . . .837 

XXVII. Morphology and Mathematics. By D'Arcy Wentwokth Thompson, 857 

XXV I II. The Poisoned Arroivs of the Abo rs and Mishmis of North- East India, 
and the Composition and Action of their Poisons. By Sir Thomas 
R. Fkaser, M.D., F.R.S., Professor of Materia Medica and Thera- 
peutics in the University of Edinburgh. (Plates XCVIII-C), . 897 



Index, /£&* m < vu $S>X .931 




TRANSACTIONS 



OF THE 



ROYAL SOCIETY OF EDINBURGH. 

VOLUME L. PART I.— SESSION 1913-14. 




CONTENTS. 

PAQH 

I. Sphxrostoma ovale (Conostoma ovale et intermedium, Williamson), a Lower Carboniferous 
Ovule from Pettycur, Fifeshire, Scotland. By Margaret J. Benson, D.Sc., Professor of 
Botany in the University of London, Head of the Department of Botany at the Royal 
Holloway College, Englefield Green, Surrey. Communicated, by Dr R. Kidston, F.R.S. 
(Plates I. and II.), ......... 1 

(Issued January 7, 1914.) 

II. Studies on the Pharmacological Action of Tetra-alkyl-ammonium Compounds. I. Hie Action 

of Tetra-methyl-ammonium Chloride. By Professor C. R. Marshall, . . .17 

(Issued December 29, 1913.) 

III. Polychxta of the family Nereidee, collected by the Scottish National Antarctic Expedition 

(1902-1904). By L. N. G. Ramsay, M.A., B.Sc, Carnegie Research Scholar, Christ's 
College, Cambridge. Communicated by Dr J. H . Ashworth. (Plate III.), . . 41 

(Issued March 30, 1914.) 

IV. On the Genus Porponia and Related Genera, Scottish National Antarctic Expedition. By 

Professor Oskar Carlgren, Universitetets Zoologiska Institution, Lund. Communicated 

by Dr W. S. Bruce. (Plate IV.), ........ 49 

(Issued March 30, 1914.) 

V. On the Fossil Flora of the Staffordshire Coal Fields. Part III.— The Fossil Floi-a of the 
Westphalian Series of the South Staffordshire Coal Field. By R. Kidston, LL.D., F.R.S. 
(Plates V.-XVI.), 73 

(Issued March 12, 1914.) 

VI. The Anatomy of a New Species of Bathydoris, and the Affinities of the Genus: Scottish National 
Antarctic Expedition. By T. J. Evans, M.A. (Oxon.), Lecturer in Zoology in the University 
of Sheffield. Communicated by Dr J. H. Ashworth. (Plates XVII. and XVIIL), . 191 

(Issued April 1, 1914.) 

VII. Rupture Stresses in Beams and Crane Hooks. By Angus R. Fulton. B.Sc, A.M.Inst.CE., 
Engineering Department, University College, Dundee. Communicated by Professor A. H. 
Gibson, D.Sc, A.M.Inst.CE., . . . .211 

(Issued April 15, 1914.) 

VIII. Scottish National Antarctic Expedition : A Description of the Systematic Anatomy of a Foetal 
\A Sea-Leopard (Stenorhynchus leptonyx), with Remarks upon the Microscopical Anatomy of 
some of the Organs. By Harold Axel Haig, M.B., B.S., M.R.C.S., late Lecturer in 
Histology and Embryology, University College, Cardiff; M'Robert Research Fellow, 
University of Aberdeen. Communicated by Professor Arthur Robinson, M.D. (Plates 
XIX.-XXIL), .225 

(Issued April 30, 1914.) 






EDINBURGH: 

PUBLISHED BY ROBERT GRANT & SON, 107 PRINCES STREET, 

AND WILLIAMS & NORGATE, 14 HENRIETTA STREET. COVENT GARDEN, LONDON. 



MDCCCCXIV. 
Price Twenty-five' Shillings and Ninepence. 



2 PROF. MARGARET J. BENSON ON 

divergence from the type under consideration than was formerly suspected — in fact, 
as the authors of the above treatise suggest, the resemblances of Sphserostoma with 
Lagenostoma are greater than with Conostoma* Hence one has been obliged to found 
a new form-genus for the geologically older type, and the term Sphserostoma was 
chosen because of the rounded form of the free part of the nucellus within which lies 
the pollen chamber. Before proceeding to describe the ovule, I wish to express my 
indebtedness to Professor I. Bayley Balfour, who presented me with the large block 
of Burntisland rock from which all but two of the slides used in the present paper have 
been obtained. 

One of these is a slide most generously put at my disposal by Dr D. H. Scott, 
F.R.S. In the course of the paper it will be shown how much is derived from this 
slide which, as a section of a Pettycur petrifaction, cannot, I think, be surpassed for 
beauty of workmanship and interest of content. I take this opportunity of expressing 
my gratitude to Dr Scott for this loan. The other slide is a fine radial section of 
the ovule, cut by Dr Gordon, a photograph of which Dr Gordon has been kind 
enough to send me (vide PI. II. fig. 8). 

Sphserostoma ovale has so far only been recorded from Pettycur deposits of the 
Calciferous Sandstone series of Scotland. In these it occurs at rare intervals, but 
generally, when found, is fairly abundant. About fifty have been sectioned from 
the above-mentioned block, but owing to the impossibility of orienting such small 
objects, a large proportion are cut obliquely. They are always closely associated with 
the vegetative organs of Heterangium Grievii, and hence the ovule has long been 
suspected of being the megasporangial apparatus of this most fern-like of all would-be 
Pteridosperms. 

Deferring the evidence which I consider places this surmise upon a surer basis, it 
will be best to describe the ovule in detail. 

II. General Features of the Organisation of the Ovule. 

(Text-figs. 1 and 2.) 

In its complete state the ovule consists of two parts — a central body representing 
l be nucellus and inner integument, and an enveloping cupule or outer integument. 

For convenience, the inner part will often be referred to as the ovule, for it is most 
common to find it bereft of its cupule. A reference to text-fig. 1, which represents a 
restoration of a median longitudinal section of both parts, will readily show their 
relationship to one another, especially if this restoration be compared with that in 
text-fig. 2, c. 

Without the cupule the ovule is 3 '5 mm. in length and about 2*2 mm. at its widest 
part. At first sight its most striking feature is the series of crests forming the so- 
called "frill" around the micropyle (text-fig. 1,/.). 

* Oliver and Salisbury, loc. cit., p. 38. 



SPH^ROSTOMA OVALE FROM PETTYCUR, FIFESHIRE. 



Within the sinus (si.), surrounded by the free part of the inner integument or 
" canopy " (ca.), may be seen the nucellar cap (n.c.) or free part of the nucellus. This 




PlaMloflL.C 



Text-fig. 1. — Diagrammatic drawing of a median longitudinal section of Spheerostoma ovale, still surrounded 
by the cupule. The description is given in the text. 



a. =archegonium. 
ab. = abscission layer. 

c. = cupule. 
ca. = canopy. 
e. s. = embryo sac. 

/.= frill. 



n.— central column surrounded by 
the pollen chamber. 
n. e. = nucellar cap. 

p. = plinth. 
p.c. = pollen chamber, 
s. —sclerotic hypoderm. 

v). = wall of the pollen chamber. 



si. = sinus. 
v.6. 1 = a vascular strand of the cupule. 
v.b. 2 = & vascular strand of the inner 

integument. 
v. 6. 3 = central supply strand to the 
ovule. 



consists of the plinth (p.) which is relatively flat, and the central nearly hemispherical 
dome or "lagenostome " which rises abruptly from the centre of the plinth. 





«*£?fts* 



** 




t-fio. 2, «, I, c, d. -Diagrammatic representations of sections across the ovule in the planes indicated in text-fig. 1. 
a, shows the eight create outside, and the eight within, the micropyle. The cupule is omitted 

''' 't th : t bv f £ e QUCellUS ^ 0lUmn=K ! a ? d r °°^ of the Pdlen Ser) standing in the centre of the 

.I.,;.;,. , ' ,S TT "^" 1,ai,t of . the inner ^tegument. Traces of the vascular bundles of the inner 

Sssi :z!: '',-;: *:;!;:,: ^m" " utcr e,mJfimiis of thc integument is ™ retot * at tMs ievcL The section is 

"' *« Lfwttfev hav^hi I, 1 ':.""" 1 " , sti,I / s,1,l " , !'" ] '" 1 by t^ cupule, c. The vascular bundles of both integuments arc inserted 

''• h °™ » J n of , be pedicel of the ovule with the single supply bundle in the centre. Around the bundle is the base 

• ' ■ l ' : ;' • '" ™nmg between thisand the epidermis is the layer regarded as the abscission layer. This 

■■ ,', f7>' 0, " n « G 7ecr,.|„ 1 . vll , thisplane. The section is represented as including the outline of a 

/• ,,„.'■ ;;'■"' r Ulc °™ fl t0 " hOTr «* "," t"™*erse section of the ovule is again octagonal at its base. 

mad. partly from a sectaon through a fallen ovule, and therefore the cupule is omitted (cf PI. II. fig. 11) 



SPH^ROSTOMA OVALE FROM PETTYCUR, FIFESHIRE. 5 

The " lagenostome " is differentiated in a manner which has so far been undescribed 
for any ovule. We find, it is true, a central column of nucellar tissue («.), surrounded 
by a lysigenic annular cavity — the pollen chamber — as in Lagenostoma and several 
other genera of Palaeozoic ovules, but Sphserostoma differs in the definite manner in 
which the roof of the pollen chamber closes the aperture made at dehiscence. As 
far as we know, the aperture was only adventitiously blocked, or the closure was due 
to growth phenomena in other seeds, but in Sphserostoma the pollen chamber is 
always found tightly closed by means of what appears to be an elastically acting 
mechanism. 

Beneath the lagenostome and plinth lies the large embryo sac (e.s.) encased in a 
framework of eight vascular bundles (text-figs. 1 and 2, c {vide b, 1)). These spring at 
the base from a single delicate strand which passes up the pedicel. In this particular 
Sphxrostoma differs from the ovules included in Messrs Oliver and Salisbury's 
groups Conostomese and Physostomese, as they are supplied by a ring of bundles. 

Accompanying the vascular bundles are large, thin-walled cells, which probably 
formed an aqueous storage tissue. They are, however, rarely preserved in the fossil 
except in the canopy. The wall of the ovule is strengthened by a layer of hypodermal 
fibres which ceases at the base and is here supplemented by deeper-lying fibres closely 
investing the single vascular strand of the pedicel, thus making the sheath complete 
when the ovule falls (PI. II. fig. 8). 

Below the sheath in the pedicel may be seen (text-figs. 1 {ab.) and 2, d) a region 
where rupture normally took place, suggesting that the cupule was left on the plant 
when the ovule was shed. 

Two series of four transverse sections through single ovules enable one to state 
definitely that the ovule is exactly circular in section in its median region, but at the 
extreme base and in the region of the canopy the transverse section gradually assumes 
an increasingly octagonal form {cf. text-fig. 2, c, with text-fig. 2, a, b, and d ; vide 
also PI. II. figs. 10 and 11). 

The outermost part of the micropyle is formed by eight lobes of the canopy, which 
bear each a large crest or lobe of the frill on the outside and a smaller one within. 
Text-fig. 2, a, is a restoration of a transverse section through this region. It is 
based on a section in slide 253, which is figured in part on PL I. {vide fig. 3). The 
individual epidermal cells constituting the crest enlarge and become hexagonal in 
section distally. They may be regarded as representing a special development of the 
epidermal cells which are characteristically developed in all the Lagenostomales, con- 
stituting the structure called the "blow-off" by Professor Oliver. 

Sphserostoma is most often found without the cupule, hence this organ is only 
represented in one of the transverse sections in text-fig. 2 {vide 2, c). 



6 PROF. MARGARET J. BENSON ON 

III. Details of Structure. 
1. The Outer Integument or Cupule. 

The cupule is probably a youth organ which did not accompany the ovule when it 
separated from the parent plant. In some sections it is represented by fragments, but 
seems to owe this preservation to the fact that parts had adhered to the slimy surface 
of the inner integument. The tissue is far more uniform than that of the inner integu- 
ment, and the vascular bundles more rounded in transverse section. 

The preservation does not admit of a decision as to the relative position of the 
phloem and xylem in each strand, nor have I been able to ascertain the number of 
the latter, but the distribution of those which can be made out is such as to suggest 
there are eight, which is also the number of the strands in the inner integument 
(text-fig. 2, c, v.b.). 

It is possible that, at the apex, the cupule was lobed like that of Lagenostoma 
Lomaxii* but most of the transverse sections passing through it show it as a continuous 
sheath (PI. I. fig. 5). 

In the specimen shown in fig. 2 of the same plate, there are at the base of the 
cupule a few longitudinally disposed fibres of cylindrical form, which are similar in 
character and position to those in the neighbouring petioles of Heterangium Grievii. 
With the exception of these and the tracheides of the vascular strands, the only tissue 
elements preserved in the cupule are parenchymatous. 

2. The Inner Integument. 

The inner integument is only free from the nucellus in the upper part, where it 
corresponds in many respects with the so-called canopy of Lagenostoma. In the 
basal three-fourths of the ovule there is no distinct layer of demarcation between 
the nucellus and the peripheral vascular part commonly regarded as the part of the 
integument adherent to the nucellus. The whole of the ovule, irrespective of the 
cupule, is surrounded by a remarkably well-differentiated epidermis, which eventually 
becomes secretory for the most part. On the exposed surface of the inner integument, 
from the pedicel up to the margin of the micropyle, the epidermal cells are longitudin- 
ally disposed, brick-shaped bodies. Each cell bears peripherally a small papilla, which 
later increases in size and eventually ruptures near the apex — a hemispherical cap being 
pushed aside by the emerging mucilage as in Lagenostoma Lomaxiif (text-fig. 3, 
a, b, and c). 

Near the micropyle the epidermal cells change their character. Outside, they 
form the structure to which I have referred as the "frill." It is seen in PI. I. fio-. ]. 
There are several sections which demonstrate that the "frill" is not continuous. In 

* Olivbb and Scott, " On the Structure of the Palneozoic seed Lagenostoma Lomaxii," Trans. Roy. Soc. 
(Lmd.), 1904 (]>. 217). 

, t (>uvi:ii and Scott, loc. cit., pi. x. fig. 28 B. 



SPH^ROSTOMA OVALE FROM PETTYCUR, FIFESHIRE. 7 

the tangential section shown in fig. 4 we see two lobes which in the specimen can 
be shown to be distinct. In an obliquely transverse section of the micropyle (fig. 3), 
which passes through the "frill" on one side, the lobe cut through is seen to be 
distinct from its neighbours, and its outline corresponds at its base with one of the 
small crests within the micropyle. Thus it can be stated that the micropyle at its 
extreme apex was surrounded by eight lobes of the canopy, each crowned by an outer 





Text-fig. 3, a. — A radial section of that part of the inner integument which lies immediately above the plane of text-fig. 2, 6, 

as plotted on text- fig. 1. 
aq. = ? aqueous tissue of the canopy ; s. = sclerotic sheath; v.l). x — the extreme apex of one of the eight 
vascular bundles that pass up the inner integument. 

b and c. — Figures to show transverse sections of young and old epidermal secretory cells, m.c. 

& = a young intact cell with papilla. Also two hypodermal fibres from the sclerotic sheath, s. 
c=an epidermal secretory cell showing liberation of hemispherical cap. 



and an inner epidermal crest, of which the outer was the larger. Lower down the 
micropyle, the epidermis contracts into a uniform layer over the eight sides of the 
octagonal sinus. Just as the sinus widens out below the micropyle, the epidermis again 
becomes papillate and secretory. The papillae are numerous and basiscopic. Below 
the surface formed by the secretory tissue lies a mass of parenchyma, behind which 
terminates the tracheal system (text-fig. 3, a, v.b.^). 

The epidermis on a level with the pollen chamber again becomes smooth and 
apparently non-secretory. Below its surface we find the large rounded cells accom- 
panying the vascular bundles throughout their entire length (text-fig. 3, aq.). They 



8 PROF. MARGARET J. BENSON ON 

.ire fairly frequently preserved in the canopy, but are rarely seen in the periphery 
of the embryo sac (slide 292, l). 

The eight regions respectively supplied by the eight vascular bundles correspond 
with those of the crests. Lower down on the same radii on which the vascular bundles 
lie, but at the very base of the sinus, between the canopy and the plinth, the epidermis 
shows some curious tongue-shaped cells which have pitted thick walls and generally 
dark brown contents (PI. I. figs. 1 and G, h,). They differ in form and appearance from 
the papillate mucilage cells, which are thin-walled and of a pale yellow colour. 

The eight vascular strands approximate closely to these cells, often obviously 
bending towards them and then retreating again above. There are probably eight 
groups rather than a continuous ring of these cells, since occasionally the radial plane 
of section does not pass through any. Such sections escape also the bundles. 

The form of these cells and their close approximation to the vascular strands 
suggest an excretory function, but their different mode of preservation suggests that 
the excretum was non-mucilaginous and hence possibly merely water. 

Let us turn now to the hypodermal structure of the exposed surface of the inner 
integument. One sees in a well-preserved specimen a uniform single layer of fibres 
approximately square in transverse section (text-fig. 3, b) and several times longer 
than broad. 

The layer appears to be less regular in some ovules — some growth in length and 
displacement probably taking place among the fibres. 

This sheath forms the only special mechanical tissue the ovule possesses, and must 
have given toughness to the coat although it was unable to afford it any great degree 
of rigidity. In several cases the ovule has been indented, or even folded like a 
collapsed bladder, before fossilisation. The sheath is not continued beneath the 
epidermis of the pedicel, but is completed by deeper-lying sclerotic cells forming a 
transverse plate, which is only perforated to allow of the entry of the single vascular 
strand into the ovule (text-fig. 1, s.). This plate forms the base of the great majority 
of the ovules examined (PI. II. figs. 8 and 11). 

3. The Abscission Layer. 

Immediately outside the sheath, as it leaves the epidermis, the tissue in the pedicel 
shows signs of breaking down. Eemains of this layer have been demonstrated in a 
transverse section of an ovule which had probably been naturally severed from the 
parent plant (PI. II. fig. 11, ab.). The tissue is seen also in the radial section of the 
ovule in slide 387 (PI. II. fig. 9, ab.) which shows the layer of separation still in situ. 
In this case the degeneration of the tissue has not yet encroached on the epidermis, 
which is still regular and continuous. 

If this be the true interpretation of this region, it indicates that the ovule normally 
fell out of the cupule, leaving that upon the parent plant. Thus it would seem probable 
that such of the ovules in Dr Scott's slide 387 as are still surrounded by a cupule had 



SPH^ROSTOMA OVALE FROM PETTYCUR, FIFESHIRE. 9 

not become at any rate normally detached. I will refer to the significance of this 
deduction later when discussing the grounds of reference of this ovule to Heterangium 
Grievii (cf. p. 12). 

4. Vascular Supply of the Ovule. 

It has already been demonstrated by an earlier reference to figs. 9 and 11 that a 
single delicate strand of tracheides enters the pedicel. This gives rise to eight strands 
which terminate behind the parenchyma abutting on the papillate epidermis near 
the micropyle (text-figs. 1, 2, and 3, a, v.b. 2 and v.b. 3 ). The constituent tracheides 
have their thickening disposed in a manner approximately spiral, and are of small 
dimensions. In several cases the strand of xylem is obviously mesarch. 

At the extreme base of the embryo sac the tracheides lie almost contiguous to one 
another, forming a sheath of which the constituent elements gradually diverge into the 
eight strands. Very frequently one or two or even three of these strands may be 
duplicated, i.e. respectively represented by two smaller, more or less laterally connected 
strands. In such cases we may meet with from nine to eleven bundles, but the position 
and size of the branching strands indicate their relationship to one another. 

5. The Nucellus and Embryo Sac. 

The nucellus exhibits features of great interest. The lower part is entirely occupied 
by the embryo sac, which never shows much contained tissue. In all the material 
investigated the embryo sac had already expanded until it almost abuts on the vascular 
strands, only a few layers of much flattened cells intervening (text-figs. 1 and 2, c, 
and PI. I. figs. 1, 5, and 6 (e.s.)). 

In the upper part the nucellus is represented by the plinth (text-fig. 1, p., and 
PL I. and II. , figs. 1,7, and 8), and by the dome-like lagenostome already referred to 
as consisting of a central column, surrounded by the annular pollen chamber. The 
plinth and dome are covered with a well-characterised but apparently non-secretory 
epidermis. 

In surface view the epidermis of the plinth is seen to consist of isodiametric 
hexagonal cells (fig. 7), and thus offers a sharp contrast to that forming the pollen 
chamber wall (fig. 10). The embryo sac has in all cases flattened or absorbed all the 
cells beneath the epidermis of the plinth. It also abuts on the base of the dome, but 
no trace of a membrane has been found beneath the excavated pollen chamber — a fact 
which probably indicates that the archegonia were formed on that part of the prothallus 
which forms the floor of the annular pollen chamber. In this position traces of them 
are occasionally found. Under the central column only a thin pellicle can be demon- 
strated, and that only occasionally. In the exact centre a thickened mass is sometimes 
seen, but it is possibly due to the destruction of the overlying cells and not to any 
special formation on the apex of the spore, as is rather suggested by its appearance. 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 1). 2 



10 PROF. MARGARET J. BENSON ON 

Directing our attention to the dome, we are very fortunately able to trace to some 
extent the development of the pollen chamber. 

6. The Structure of the Pollen Chamber. 

In the older ovules there is a clear broad space seen in radial section on either 
side of the central column and roofed over by a single layer of epidermal cells. 
In longitudinal sections it will be observed that the column consists of regularly 
arranged tiers of cells. In this latter feature Sphserostoma resembles Lag eno stoma . 

Tn transverse section the form of dome and column is found to be circular. 

In the nearly radial section of a young ovule, shown in PI. I. figs. 1 and 2, the 
tissue in the annular region, to be later replaced by the pollen chamber, is seen to be 
undergoing lysigenic degeneration. A difference between the younger and older domes 
is seen also in the character of their epidermis. In the younger, the epidermis is 
composed of thin-walled cells with flat peripheral walls. In the older, the epidermis 
over the pollen chamber has entirely changed its character, while that over the column 
has remained unchanged. Each cell over the pollen chamber has its walls sharply 
differentiated, i.e. the outer and basal walls remain thin, and all the four walls vertical 
to the surface are thickened in such a way that they appear of an opaque dark brown 
colour, while the outer and basal walls are quite transparent. Thus the whole roof or 
wall of the pollen chamber has much the appearance of a multiseriate annulus of such 
a sporange as that of Senftenbergia. The resemblance is heightened by the obliquity 
of the end walls of the cells (PI. II. figs. 7 and 10). 

The circumscissile dehiscene, by means of which presumably the pollen grains entered 
the chamber, is effected in a very definite manner. The central column becomes severed 
entirely from the roof of the pollen chamber along a line just within the margin of the 
epidermis of the column. 

Thus marginal epidermal cells of the column remain attached to the roof-cells 
(vide PI. 1. fig. 6). There is no loss of surface cells, the dehiscence is schizogenic, and in 
some respects the whole apparatus is curiously reminiscent of the peristome in the 
sporogonium of Polytrichum, which opens around a persistent, circular diaphragm. 
The mechanism, however, of the closure in the two cases must have been very 
different, and the resemblance between them merely adventitious. 

There are tetrahedral spores found in the pollen chambers of seeds in slides 217, 
241, 270*1, and 304 - 4. Occasionally they occur still arranged as tetrads. These spores, 
which are presumably the pollen grains, are from 27-29m in their widest dimension, 
and thus much smaller than those * attributed to Lyginodendron, which are from 50- 
70/j- in diameter. 

Traces of a species of microsynangium do, however, occur with the Sphwrostoma, 
and its spores resemble in size and form those in the pollen chamber. Description of 

* Kiuston, "On the Microsporangia of the Pteridospennece," Trans. Boy. Soc, 1906, vol. cxcviii. p. 423. 



SPH^EROSTOMA OVALE FROM PETTYCUR, FIFESHIRE. 11 

this body must be deferred until another occasion. So also must the description of 
some curious bodies which often accompany the ungerminated grains in the pollen 
chamber. 

No satisfactory specimen has yet come to hand in which the pollen chamber is 
open. One of Williamson's slides, preserved in his collection at the British Museum 
at South Kensington, and figured in his Eighth Memoir,* suggests this condition, but 
the appearance may be due to injury before fossilisation, as only one side is open. 

7. The Processes assumed to occur during the Opening and Closing of the 

Pollen Chamber. 

That the important function of securing and nursing the pollen was correlated with 
a series of progressive changes cannot be doubted. When the excavation of the pollen 
chamber was complete, the epidermis of the nucellar cap underwent circumscissile 
dehiscence along a line just within the periphery of the persistent column of tissue in 
the centre. The roof of the chamber was thus made to overlap the column when it 
returned, after dehiscence, to its original position (cf. figs. L = state before dehiscence 
and 6 = state after closure). In fact, the relation of the margin of the column to the 
roof of the pollen chamber resembles that of the rebate of a box to its lid. Hence the 
downward curvature of the roof is prevented from being so excessive as to obliterate 
the pollen chamber. 

The growth in length of the roof-cells and the deposition of thickening on their 
vertical longitudinal walls would necessarily have set up a centrifugal strain tending 
to bring about the circumscissile dehiscence. This was further aided by the mucilagin- 
ous degeneration of the subjacent tissue. 

After dehiscence the roof would straighten elastically and thus an annular stomium 
would be formed. Passage to the lower part of the sinus would be at least partially 
blocked because the liberated margin of the roof of the pollen chamber would nearly 
abut on the basiscopic papillae around the upper part of the sinus. Pollen would thus 
be prevented from straying into the sinus. 

The mucilage glands would meanwhile have come into play, thus completing the 
preparation for pollen. If grains were now to enter the micropyle they would be 
caught in the freshly exuded watery mucilage and be drawn with it into the pollen 
chamber. The entry must have been aided by the subsequent downward curvature of 
the roof of the chamber. 

This return to the former position may have been due to an increase in their 
turgescence on the part of the roof-cells. This is suggested by the domed form of the 
peripheral cell-walls of the epidermis covering the older pollen chambers. These 
peripheral walls offer a marked contrast not only to the thick vertical cell- walls, but 
even in this respect to the basal walls, which, although thin and transparent, 
are flat. 

* Williamson, he. cit., pi. xii. fig. 83. 



12 PROF. MARGARET J. BENSON ON 

The tongue -shaped cells already described, which otherwise seem so enigmatic, 
receive their interpretation if they were the source of the water required for the closure 
of the pollen chamber. 

Summary of Stages. 

1. Growth in length and special thickening of the roof-cells of the pollen chamber. 
Concomitant lysigenic degeneration of the subjacent tissue leading to the excavation 
of the pollen chamber. 

2. Circumscissile dehiscence and consequent formation of a stomium by the upward 
movement of the free margin of the roof. 

3. The retention of the pollen grains on the downward curvature of the roof, 
which thus returns again to its original position and closes the pollen chamber. 

8. Diagnosis of Sphxrostoma, n.g. 

The ovule resembles those of the Lagenostoma series in its general organisation, 
e.g. the nucellus is surrounded by an inner integument of radial symmetry and appar- 
ently multiple origin, and also by an outer integument or cupule. The inner integu- 
ment is only free around the pollen chamber, but the cupule is free from the base 
upwards. Both integuments have a multiple vascular supply — the strands of the inner 
integument taking their origin from the single central bundle of the pedicel. Sphwro- 
stoma is distinguished from the ovules of the Lagenostoma series by the following 
characters : — 

1. The whorl of epidermal crests around the micropyle. 

2. The nearly hemispherical form of the lagenostome which rises from a somewhat 
flat plinth. 

3. A pollen chamber relatively wider than that of Lagenostoma. 

4. The persistence of the epidermis over the central column of the lagenostome. 

5. A skeletal structure of less robust charater, with an indication of aqueous tissue 
in the region of the canopy. 

Dimensions. — The ovule is a little under 3 '5 mm. in length without the cupule 
and 2*2 mm. at its widest part. 

Horizon and Locality. — Sphterostoma ovale has so far only been recorded from 
the Calciferous Sandstone Series of the Lower Carboniferous rocks of Pettycur, 
Kifeshire, Scotland. 

[V. The Grounds for the Provisional Reference of the Ovule Sph^erostoma 

OVALE TO HETERANGIUM GrIEVII. 

As already mentioned, this ovule has long been surmised to be the mega- 
sporangial apparatus of LLeterangium Grievii. The association between the two is 
of ;i very constant and striking nature. For example, there are some score of plants 



SPHSEROSTOMA OVALE FROM PETTYCUR, FIFESHIRE. 13 

of Heterangium Grievii in the same block with the ovules here described. Many of 
the plants are exceptionally well developed, and three are giving off adventitious roots. 
In the course of three years, during which time I have had hundreds of slides from 
this block under observation, I have not been able to detect a trace of any other 
plant the nature of whose sporangia is not known, except two undescribed stems 
apparently belonging to immature plants or other species of Heterangium. 

Hence the fact that numerous ovules lie among the petioles and rooted stems of 
Heterangium Grievii is undoubtedly suggestive of actual reference. 

The argument from association is supplemented by internal evidence. The ovule 
shows an unmistakable resemblance to Lage?wstoma, which is acknowledged to be 
the type of ovule borne by Lyginodendron. 

But Lyginodendron, both in the external morphology of its vegetative organs and 
in their anatomy has long been regarded as showing close affinity with Heterangium. 
The resemblance between their megasporangial members would be similarly great if 
Sphserostoma ovale belonged to Heterangium. Such differences as do obtain between 
the two ovules are such as would be expected between an older and a later type. 
In Sphserostoma the canopy shows less complete integration (i.e. the crests are free) 
and the mechanical tissue is less developed than in Lagenostoma. The method of 
dehiscence of the nucellus is more fern-like and the pollen grains are smaller. These 
differences are all consistent with the view that we are dealing with a more primitive 
ovule in Sphserostoma than in Lagenostoma, but with one in which the ground-plan 
of the Lyginodendron ovule had been laid down. 

This evidence from comparative considerations receives important corroboration 
from certain features of Dr Scott's slide, C.N., 387, already referred to. 

Many of the ovules are still surrounded with an intact cupule and hence, if the 
interpretation of the tissue in the pedicel as an abscission layer be correct, must have 
been still attached to the frond, or part of the plant on which they grew. 

To investigate this matter the three longitudinal sections which were nearest the 
median plane were examined, with the following results : — 

a was found to be exactly radial at its base ; 

b was radial at the micropyle but slightly tangential at the base ; 

c was in the plane which passed through the plinth but just escaped the pollen 
chamber. At the base it also just escapes the pedicel of the ovule. 

Sections a, b, and c were found in each case to be accompanied by a petiole in the 
expected position and cut in a plane exactly corresponding to that of the ovule. Thus 
a shows a radially cut petiole, b shows one in which the vascular bundle is just touched 
upon, and c shows one with only the cortex occurring. For the case referred to as 
a, reference should be made to PL II. fig. 9. 

As already pointed out in an earlier part of the paper, most of the ovules show no 
cupule. Such ovules in no single case, although they are far more frequently met with, 
show similarly disposed petioles of Heterangium. 



14 PROF. MARGARET J. BENSON ON 

To conclude, the evidence in support of the reference of Sphserostovna ovale to 
Heterangium Grievii may be summed up as follows : — 

1. Association. 

2. Comparative morphology, i.e. resemblance to Lagenostoma. 

3. Strong suggestions of continuity in the only three cases in which sections have 
been obtained in a suitable plane, of ovules judged by independent evidence to have 
been still attached to the parent plant. 

In conclusion, I wish to bear testimony to the steady and able help I have had 
from my laboratory attendant, C. H. Wells, who has made all but two of the 
preparations used in this paper. 



V. EXPLANATION OF PLATES. 

S. = Scott Collection. 
R.H.C. = Royal Holloway College Collection. 

Lettering of Figure*. 



a. = archegonium. 
ab. = abscission layer. 

c. = cupule. 
ca. = canopy. 
e.s. = embryo sac. 

/. = frill. 

h. = tongue-sbaped cells at the base of sinus. 
m. = micropyle. 
m.e. = mucilage epidermis (Prof. Oliver's "blow-off") 

n. = persistent column of nucellar tissue. 



p. = plinth. 
p.c. = pollen chamber. 
pet. = petiole. 

s. = sclerotic hypodermal sheath. 

si. = sinus. 
•y.6 n = vascular bundle of inner integument. 
v.b. 2 = vascular bundle of cupule. 
v.b. 3 = vascular bundle of the pedicel of the ovule. 

to. = wall of the pollen chamber. 



Plate I. Figs. 1-6. 

Fig. 1. Radial section of the upper part of a young specimen of Sphserostoma ovale enclosed in a 
cupule (a). The lagenostome shows an early phase in the development of the pollen chamber (>.e. excava- 
tion incomplete). The tongue-like cells (h.) are formed, but the mucilage papilla? are not yet developed on 
the inner surface of the canopy. Those on the outer surface are not yet dehisced (m.e.). The embryo sac 
(e.s.) shows traces of an archegonium (a.). Supporting the crests are seen horn-like prolongations of the 
sclerotic sheath (s.). 

S., C.N., 387x70xf. 

Fig. 2. Drawing from a photograph of the whole section of a young ovule, part of which is represented 
in fig. 1. 

S., C.N,, 387 x about 30 x f. 

Fig. 3. Part of an obliquely transverse section through the micropyle in the plane plotted on fig. 1. 
It shows the corresponding crests on the inner and outer surfaces. At this level the sclerotic sheath is 
interrupted between the pairs of crests. 

R.H.C., C.N., 253 x about 70 x ^ ! . 



SPH^EROSTOMA OVALE FROM PETTYCUR, FIFESHIRE. 15 

Fig. 4. A tangential section through the upper part of a similar ovule, showing two crests of the " frill." 

S., C.N., 387 x about 80 x f . 

Fig. 5. Drawing from a photograph of part of a transverse section of an ovule still enclosed in the 
cupule. The vascular bundles of cupule and inner integument can be seen, also the embryo sac and mucilage 
epidermis (in.e.). 

S., C.N., 387 x about 60 x f . 

Fig. 6. A slightly tangential section of the upper part of an ovule, showing a pollen chamber after 
dehiscence. The column (n.) is well preserved, and shows the ledge referred to in the text as a " rebate." 

R.H.C., C.N., 241 x 70 x f . 

Plate II. Figs. 7-11. 

Fig. 7. Drawing from a photograph of an oblique section through an ovule to show the distribution of 
the eight vascular bundles of the inner integument. This section also exhibits the surface cells of the plinth 
and column, and the contrast they afford to those of the pollen chamber wall (w.). 

R.H.C., C.N., 287 x about 70 x f 

Fig. 8. A radial section through the ovule, enlarged from a negative kindly lent for the purposes of this 
paper by Dr Gordon. The tissue (c.) outside the inner integument is probably cupular. 

x about 27 x f . 

Fig. 9. Drawing from a radial section of a somewhat crushed young ovule. At + the epidermis of the 
radially cut petiole of Heterangium Grievii is wrinkled and appears continuous with the tissue of the cupule, 
but the cell-walls are not well preserved. The cupule is obviously continuous with the ovule by its epidermal 
cells on the left of the section. The pedicel bundle can be traced down the centre and is seen to give rise 
to the inner integument bundles. The section suggests that the ovule grew terminally on the petiole. 

S., C.N., 387 x 62. . 

Fig. 10. Part of a transverse section at the level of the roof of the pollen chamber, showing it in surface 
view. The integument is octagonal, and the vascular bundles are well developed at this level. 

R.H.C., C.N., 290'3 x about 70 x f . 

Fig. 11. A transverse, slightly oblique section across the base of an ovule after it has fallen. In the 
centre can be seen the delicate strand of tracheides which perforate the basal sheath of sclerotic cells. The 
disorganised cells immediately outside this sheath to the left may possibly represent remains of the abscission 
layer shown in fig. 9 (from an ovule probably still attached to a petiole). On the right may be seen the 
sclerotic sheath cut tangentially and clearly indicating the octagonal form of the base of the ovule. The 
mucilage epidermis is shown in a condition very characteristic of the shed ovule. The cells have become 
so swollen that they are detached from the hypoderm and form a pale-yellow, halo-like area all round 
the section. 

R.H.C., U.N., 246 x 87 x f . 



Trans. Roy. Soc. Edin r 



Benson : Sph^erostoma ovale. 



Vol. L.— Plate I. 






-.i.2 





vf^'W/P 



s\ n 




EJ 



M'Farlaiic * ErtWne. Uth.. Edin. 



Trans. Roy. Soc. Edin r 



Benson : Sph^erostoma ovale. 



Vol. L.— Plate II. 





U.S. 



^J# 9 





v.b.3 




M'Farlute & Enkiac, Lith.. KJin. 



( 17 ) 



II. — Studies on the Pharmacological Action of Tetra-Alkyl-Ammonium 
Compounds. By Professor 0. R. Marshall. 

(MS. received April 30, 1913. Read November 17, 1913. Issued separately December 29, 1913.) 

I. THE ACTION OF TETRA-METHYL-AMMONIUM CHLORIDE. 

In a paper on " The Pharmacological Action of Protocatechyl-tropeine," communi- 
cated to the Society in 1909, # I drew attention to the fact that this substance, when 
injected intravenously in certain doses produces transient paralysis of the respira- 
tion ; and I mentioned further that Tappeiner t had described a similar temporary 
cessation of the respiration after the intravenous injection of certain quaternary 
isoxazol and pyrazol compounds, and of tetra-methyl-ammonium chloride, and Pohl,} 
after the intravenous injection of some quaternary papaverine derivatives. Tappeiner 
came to the conclusion that the effect was due to stimulation of the terminations 
of the fifth cranial nerve in the nose ; that it was, in fact, of the nature of a 
Kratschmer-Hering reflex, since he was unable, in the case of methyl-phenyl- 
isoxazol-methochloride, to produce cessation of the respiration after anaesthetising 
the nasal mucous membrane with cocaine ; and Iodlbauer,§ working in Tappeiner's 
laboratory, also found that ansesthetisation of the nasal mucous membrane prevented 
the cessation of the respiration produced by tetra-methyl-ammonium chloride. Pohl, 
on the other hand, was able to produce this temporary paralysis of the respiration 
after section of the ophthalmic branches of both fifth nerves, and consequently he 
concluded that the effect was due to an action on the respiratory centre. I came 
to the same conclusion, since the effect was still obtained with protocatechyl-tropeine 
after section of both fifth nerves in the base of the skull and after section of both 
phrenic nerves, and was not synchronous with the effect on the circulation or with 
the paresis of the nerve-endings in the muscles of the hind limbs. Further work 
with tetra-methyl-ammonium chloride, however — my stock of protocatechyl-tropeine 
being exhausted, — showed that the effect was in large measure peripheral and due 
to a transient paresis of the nerve-endings in the respiratory muscles. 

Action on Motor Nerve- Endings. 

The effect of tetra-methyl-ammonium compounds on the motor nerves was first 
observed by Crum Brown and Fraser.|| They showed that the iodide when injected 
into frogs produced muscular paralysis due to an action on the myo-neural junctions ; 

* Trans., xlvii., pt. ii. p. 273 [1909-10]. t Arch.f. exp. Path. u. Pharm., xxxvii. p. 325 [1896], 

% Arch. Internat. de Pharmacod., xiii. p. 479 [1904]. § Arch. Internat. de Pharmacod., vii. p. 183 [1900]. 
|| Proc. Boy. Soc. Edin., vi. p. 556 [1869]. 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 2). 3 



18 PROFESSOR C. R. MARSHALL ON THE 

and their observations were corroborated by Rabuteau * and Brunton and Cash.1" 
Later, Dufaux,| Santesson and Koraen,§ and Iodlbauer || showed that the same effect 
was obtained with tetra-methyl-ammonium chloride. 

So far as I am aware, no experiments demonstrating this action on mammals 
have been described. Rabuteau,H in the two experiments he made with tetra-methyl- 
ammonium iodide on dogs, observed muscular paralysis as one of the symptoms ; and 
Brunton and Cash,** after injecting 1 g. of the same compound into a rabbit, found 
the animal lying apparently paralysed two minutes after the administration. When 
half this dose was injected, they noticed the head falling to one side two and a half 
minutes afterwards, but one and a half minutes later violent movements of the limbs 
occurred. IoDLBAUER,tt working with the chloride, had similar experiences. The 
animal usually fell on its side, and there were frequently convulsive movements and 
later fibrillary tremors. Jacobj and Hagenberg|| also noticed muscular paralysis 
after tetra-methyl-ammonium tri-iodide. But none of these observers appears to 
have investigated the cause of the paralytic symptoms. 

My own experiments have been made on rabbits and cats during a state of 
anaesthesia or after excision of the cerebral hemispheres. The rabbits were anaesthetised 
with ether, or, in the case of animals to be decerebrated, with a mixture of chloroform 
and ether ; the cats with chloroform followed by ether. The blood-pressure and 
respiration were recorded, and one of the nerves supplying a fore or hind limb was 
isolated and stimulated continually during an injection or intermittently before and 
after an injection. The contraction of the limb was recorded by inserting a hook 
into the limb and connecting this by thread working over pulleys to a writing lever. 
This method of recording; the contractions of the limb was found better for the 
purpose in view than that of recording the contractions of individual muscles, 
probably owing to less interference with the blood supply. The injection of the 
substance was made in the case of rabbits into the right anterior facial vein and 
in the case of cats into the right external jugular vein. 

Both in rabbits and cats the intravenous injection of 0'5 to 1 mg. p. kg. body- 
weight diminishes the excitability and often paralyses the motor nerve-endings, the 
muscles themselves still retaining their irritability, over the whole of the body. 
Rabbits seem to be somewhat more susceptible than cats, but otherwise the effects 
on the two animals are identical. When the injections are made into the facial or 
external jugular vein, the muscles of the fore part of the body are earliest and most 
markedly affected. After a dose of 1 mg. p. kg. moderately quickly injected, the 
nerve endings in these muscles are more or less paralysed within ten seconds of the 
commencement of the injection, and in susceptible animals may remain paralysed for 
several minutes. Fairly quick recovery then occurs. 

* ComjA. rend., lxxvi. p. 887 [1873]. t Trans. Roy. Soc. Lond., clxxv. p. 215 [1884]. 

X Dissert. Berlin., 1888, quoted by Santesson and Koraen, p. 220. 

§ Skand. Arch./. Physiol., x. p. 220 [1900]. || Loc. cit., p. 185. IT Loc. cit. 

** Loc. cit., p. 209. tt Loc. cit., p. 189. %% Arch.f. exp. Path. u. Pharm., xlviii. p. 48 [1902]. 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL- AMMONIUM COMPOUNDS. 



19 




20 



PROFESSOR C. R. MARSHALL ON THE 



The action on the fore limbs of a cat is shown in the uppermost tracings (L) 
of figs. 1 and 2. Both figures are taken from the same experiment. Previous to 
the injection shown in fig. 1 an injection of half the dose of tetra-methyl-ammonium 
chloride had been given. The first figure shows the results of intermittent stimula- 
tion of the musculo-cutaneous nerve with the secondary coil at 40 cm., the primary 



L 
R 




H 

b 
t 



Fig. 2. — Effect of tetra methyl-ammonium chloride on contraction of fore limb produced by continuous stimulation 
of musculo-cutaneous nerve. Same animal as fig. 1. Letters as in previous figure, x J. 

current being obtained from an accumulator with an B.M.F. of two volts. The 
wavy appearance of the earlier part of the tracing is due to the presence of small 
spontaneous rhythmical contractions of the limb. Almost immediately after the cessa- 
tion of the injection these spontaneous movements ceased and stimulation of the nerve 
a few seconds later produced a scarcely appreciable effect. In this case total paralysis 
to this strength of stimulus lasted six minutes, then recovery gradually occurred ; but 
the nerve did not regain its former irritability, and the stimulus was consequently 
increased. 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL- AMMONIUM COMPOUNDS. 21 

The effect of stimulation with the secondary coil at 30 cm. is shown in fig. 2, 
which also shows the action of the same dose of tetra-methyl-ammonium chloride 
during continuous stimulation of the nerve. The cessation of the contraction seven 
seconds after the commencement of the injection is striking. The method of con- 
tinuous stimulation does not, however, show recovery from the effects of the drug 
owing to fatigue being so easily induced. For this purpose intermittent stimulation 
is necessary. 

When strong or moderately strong intermittent stimuli are employed, complete 
paralysis of the nerve-endings may not be observed after the injection of 1 mg. p. kg. 
body-weight of tetra-methyl-ammonium chloride. Each stimulation after the admini- 




Fig. 3. — Contractions of fore limb of rabbit produced by stimulation of musculo-cutaneous nerve during recovery 
from tetra-methyl-ammonium chloride. For description see text. 

stration of the drug may produce a well-marked, although less than normal, contraction ; 
but the contraction is not maintained, and notwithstanding the continuance of the 
stimulus, quickly falls to or near the abscissa (fig. 3). This contraction, indeed, may 
be so short as to resemble a simple twitch. This effect of apparent rapid exhaustion was 
first shown by Boehm * in the case of curarin. 

Occasionally another effect — a staircase effect — is obtained by stimulating a nerve 
during recovery from the paralysing action of tetra-methyl-ammonium chloride, and 
this, combined with the rapid exhaustion of the nerve -endings, produces a curious, 
humped-back form of curve. This is shown in fig. 3. The animal, a rabbit, was 
anaesthetised with ether; the brain was excised at 10.43, and the anaesthetic discon- 
tinued at 11.0. The musculo-cutaneous nerve was isolated and divided, and the 

* Arch. f. exp. Path. u. Pharm., xxxv. p. 16 [1894]. 



22 PROFESSOR C. R. MARSHALL ON THE 

movements of the limb were registered as in the previous experiment. Stimulation 
with the secondary coil at 26 cm. produced a well-marked contraction, sometimes with 
a slight staircase phenomenon. An injection of 1 mg. tetra-methyl-ammonium chloride 
was made into the right facial vein at 12.3, and after preliminary convulsive contrac- 
tions, produced paresis of the nerve-endings. Stimulation of the nerve at 12.7 with 
the secondary coil at 26 cm. produced a simple muscle twitch. At 12.11, with the 
coil at 22 cm., a slight sustained contraction was obtained, and at 12.19, with the 
coil at 20 cm., the divided humped-back contraction shown at (a). The effect is 
shown better at (b), the curve obtained by continuous stimulation of the nerve with 
the same strength of stimulus at 12.21. 

The cause of the staircase phenomenon in tetra-methyl-ammonium chloride 
poisoning has not been further investigated. It differs from that produced under 
physiological conditions by adjusted repetitions of a stimulus in being preceded by a 
well-marked rapid contraction which, in some cases, exceeds in height the summit of 
the staircase, thus resembling in form the initial contraction and reflex rebound con- 
traction observed on stimulating an uncut mixed nerve with a single induction shock* 
and in the muscle requiring a much longer period of rest before it can be reinvoked. 
As is seen in (a), a repetition of the stimulus fifteen seconds after the summit of the 
staircase has been reached, is followed by no staircase phenomenon, and this is also true 
if the stimuli are repeated at half-minute intervals. In this case the only observable 
difference shown by each successive curve from that of the latter half of (a) is a more 
rapid exhaustion. It is not until an interval of one to one and a half minutes between 
the successive stimulations is allowed that the staircase effect begins to reappear. 

In one experiment the spinal accessory was isolated and stimulated intermittently, 
along with the phrenic nerve, and the contractions of the sterno-mastoid muscle recorded 
before and after injection of tetra-methyl-ammonium chloride. The effects obtained 
were so similar to those observed in the case of the fore limb that further description is 
unnecessary. 

In the case of a hind limb the effects observed after the injection of tetra-methyl- 
ammonium chloride from stimulation of the sciatic or crural nerve are similar to those 
occurring in a fore limb; but when the injections are made into the external jugular 
vein the effects are less marked, and are later in appearing. Thus in a rabbit an- 
aesthetised with ether the injection of 1 mg. tetra-methyl-ammonium chloride into the 
external facial vein merely caused gradual diminution in the contractions of the hind 
limb, produced by stimulation of the sciatic nerve, for two minutes, at the end of 
which time only a slight twitch was obtained from continued stimulation. Eecovery 
commenced one and a half minutes later, and was nearly complete six minutes after the 
injection. The respiration, on the other hand, rapidly diminished in extent, and had 
almost ceased eleven seconds after the commencement of the injection. It began to 
increase ten seconds later, and was normal a minute after the beginning of the injection. 

*^Cf. Sherrington, Journ. of Physiol., xxxvi. p. 202 [1907]. 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL-AMMONIUM COMPOUNDS. 23 

Similarly, when the injection of tetra-methyl-ammonium chloride was made during 
stimulation of the sciatic nerve, the fall of the muscle lever was gradual, and it did not 
reach the abscissa before the respiration had recovered from its paralysis. The same 
want of synchronism between the respiratory paralysis and the paralysis of the hind 
limbs occurred after the injection of protocatechyl-tropeine, and led me to minimise the 
peripheral origin of the respiratory paralysis. 

The effect of tetra-methyl-ammonium chloride on the terminations of the phrenic 
nerve was determined in a number of instances, and found to be similar to that on the 
nerves of the fore limbs. It will be referred to in considering the action of this 
substance on the respiration. 

Effect on Decerebrate Animals. — After division of the brain through the quadrate 
bodies and crura and delay for half to three-quarters of an hour after cessation of the 
ansesthetic to allow of its excretion, the intravenous injection of tetra-methyl-ammonium 
chloride produces in cats, less commonly in rabbits, immediate short clonic convulsions 
(fig. 4). These convulsions may terminate in death. If not, the convulsions cease or 
almost cease, and more or less paralysis of the motor nerve-endings may be demonstrated. 
Even if convulsions do not occur, increased excitability can be shown to be present for 
a few seconds after the injection by blowing on the animal. The reflex response to 
this form of excitation is markedly increased immediately after the injection, and is 
still present when the respiration has ceased (fig. 5). It quickly diminishes, however, 
and a few seconds later no response can be elicited. The slight normal response returns 
with or soon after the recommencement of the breathing, but no distinct increased 
excitability occurred in the experiments in which this form of stimulation was tried 
until another injection was made. Occasionally slight convulsive movements were 
noticed in ansesthetised animals after a second or later injection. 

Most observers who have investigated the general effects of tetra-methyl-ammonium 
compounds on mammals mention convulsions as a prominent symptom. Brunton and 
Cash,* for example, state that after the injection hypodermically of 0*2 g. tetra-methyl- 
ammonium iodide to a rat, the animal was affected at once with powerful convulsions ; 
and similar results with much smaller doses of the chloride on mice and guinea-pigs 
were observed by IoDLBAUER.f 

The seat of origin of the convulsions has not been determined. The simple response 
to external stimulation (fig. 5) points to the spinal cord being involved ; but as the 
convulsions exhibit a similar form to those produced by drugs acting on the basal 
ganglia, it is probable that these structures are also affected, and as such drugs also act 
on the motor areas of the cerebrum, these centres may be influenced as well. 

Effect on Frogs. — In the frog convulsions are not produced by the injection of 
tetra-methyl-ammonium chloride into a lymph sac. The predominant symptom in 
these animals is muscular paresis or paralysis. After a minimal paralytic dose the 
respiratory movements cease within two minutes, and a minute later the animal is 

* Loc. cit., p. 207. + Loc. cit., p. 190. 



24 



PROFESSOR C. R. MARSHALL ON THE 



unable to turn over when laid upon its back. The effect is upon the nerve-endings 
mainly as may be shown by electrical stimulation of the nerves and muscles after 
the intoxication. 

According to Iodlbauer* the minimal paralytic dose for the edible frog is 0*005 mg. 




Fig. 4. — Clonic spasms of fore limb 
of decerebrate cat produced by 
2 mg. tetra-metliyl-amnionium 
chloride intravenously forty- 
five minutes after excision of 
cerebrum and cessation of anaes- 
thetic. Injection shown at X. 
Time in seconds. 



H 

b 

t 




Fig. 5. — Reflex excitability during paralysis of respiration in decerebrate rabbit after 
intravenous injection of 2 mg. tetra-methyl-ammonium chloride. Rabbit. 
2200 g. Cerebrum excised 2.50. Both vagi cut. Atropine 5 mg. injected at 
4.4. At points marked X animal was sharply blown upon. Letters as in 
previous figures. 



p. g. body-weight; according to Santesson and Koraent 0"01 mg. p. g. body-weight. 
The latter observers give as the minimal paralytic dose for the grass frog 0'12 mg. p. g. 
body-weight. My own experiments with smaller frogs indicate that it is 0-08-0"09 mg. 
p. g. body-weight. 

Dufaux| and Iodlbauer § describe fibrillary contractions as occurring in frogs 

* Loc. cit., p. 186. t Loc. cit., pp. 221, 222. J Loc. cit. § Loc cit., p. 188. 



PHARMACOLOGICAL ACTION OF TETKA-ALKYL-AMMONIUM COMPOUNDS. 25 

after tetra-raethyl-ammonium chloride. They do not appear to have been noticed by 
Santesson and Koraen,* and they were not observed in my experiments with winter 
frogs. On the contrary, I found that this substance would inhibit or prevent the 
fibrillary contractions produced by tetra-methyl-ammonium chloride. 

They occurred, however, to a slight extent when an isolated muscle was steeped in 
certain concentrations of tetra-methyl-ammonium chloride, and I have frequently seen 
fibrillary twitchings in rabbits occurring both spontaneously and as the result of 
stimulating a nerve. To this point I shall return in a later communication. 




vj\rumruTj\nnnr\j\rL r \riJ\^ 



Fig. 6. — Effect of 1 ing. tetra-methyl-ammonium chloride intravenously on respiration and blood-pressure of 
etherised rabbit. Weight 1650 g. Letters as in previous figures. 



Effect on Respiration. 

In anaesthetised animals the most interesting effect on the respiration is the 
temporary paralysis which follows the intravenous injection of certain doses of this 
drug. The effect is most typically seen in rabbits, and is shown in fig. 6. The effect 
of a larger dose on a cat is seen in fig. 1. As will be observed, the respiration ceases 
or almost ceases a few seconds after the injection of the substance, and some seconds 
later recommences, at first slightly and somewhat less frequently than normally, and 
then with increasing depth and frequency until the normal is reached. The dose 
required to produce the effect is 0'5 to 1 mg. p. kg. body- weight intravenously, and it 
is necessary to make the injection moderately quickly (in about five seconds) in order to 
obtain for a sufficiently brief interval the proper concentration of the substance in the 

* hoc. cit., p. 220. 
TRANS. ROY. SOC. EDIN, VOL. L. PART I. (NO. 2). 4 



26 PROFESSOR C. R. MARSHALL ON THE 

blood. If the injection is made much more slowly, greater or less retardation and 
shallowness, but not actual stoppage, of the respiration may be produced. If larger 
doses are employed and rapidly injected, the respiration is quickly and completely 
paralysed, and it usually fails to restart spontaneously ; or, if it does commence again, 
it remains inefficient. In such cases artificial respiration is generally able to reinstate 
the respiration unless the dose administered has been excessive. 

Effect on Decerebrate Animals. — On decerebrate animals, if time has been allowed 
for the influence of the angesthetic to pass away, the effect of tetra-methyl-ammonium 
chloride on the respiration is, in some respects, different from that described as 
occurring in anaesthetised animals. In the latter no preliminary stimulation of the 
respiration, beyond a slight increase in depth of one or two respirations, occurs with 
any dose of the drug ; in the former this effect is a common feature of its action. And 
not only may there be the appearance of stimulation before the paralysis occurs, but 
also the recovery from the paralysis is quicker and is often accompanied by greater 
respiratory activity. Further, in some decerebrate animals the quantity of tetra- 
methyl-ammonium chloride necessary to cause respiratory paralysis has been several 
times that required for anaesthetised animals ; and in these animals especially the 
stimulant effect on the respiration has been very marked. If, however, the decerebrate 
animal is lightly anaesthetised, the effects are similar to those seen in anaesthetised but 
otherwise normal animals. 

The stimulating effect on the respiration of a decerebrate animal of a dose of tetra- 
methyl-ammonium chloride usually sufficient to paralyse the respiration of an anaes- 
thetised animal, and the paralysing effect of a large dose, are shown in fig. 7. The 
animal, a rabbit (2450 g.), was anaesthetised with ether containing a little chloroform. 
At 11.3 the brain above the corpora quadrigemina was removed, and between 11.10 
and 11.20 the fifth cranial nerves were cut proximal to the Gasserian ganglia. The 
anaesthetic was then stopped. At 11.36, 0*6 mg. tetra-methyl-ammonium chloride 
was injected into the right facial vein, with much the same result as is shown in the 
first part of the figure. At 11.44^, 1 mg. was injected, and, as is seen, the respiration 
was powerfully stimulated. No depression or paralysis occurred. At 11.58 the same 
dose was injected more slowly, and a similar but less marked effect resulted. Before 
the effects of this dose had passed away, 2 mg. was injected, and further transient 
stimulation followed by temporary paralysis obtained. Subsequently 5 mg. was injected, 
with the result shown in the second part of the figure. Occasional shallow respiratory 
movements occurred later, but the respiration failed to re-establish itself and the 
animal succumbed. 

As the effects obtained on decerebrate animals were slightly variable, and as the 
matter is of some importance for the purposes of this paper, a summary of twelve 
experiments made on rabbits is given. The animals were anaesthetised with ether, and 
the anaesthesia maintained with a mixture of ether and chloroform until the mid-brain 
was divided and the cerebrum removed. In some experiments the fifth cranial nerves 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL-AMMONIUM COMPOUNDS. 



27 



were divided close to the Gasserian ganglia ; in one experiment the ganglia were 
removed. The anaesthetic was then stopped. Cannulse were introduced into the right 
common carotid artery and the right anterior facial vein, for the purposes of recording 
the blood-pressure and injecting the substance respectively. The respiration was 
recorded by attaching a phrenograph or one limb of the tracheal cannula to a recording 
tambour. The time occupied by the injections, unless otherwise stated, was about 
five seconds. 








Fig. 7. — Effect of tetra-methyl -ammonium chloride on respiration and circulation of decerebrate rabbit. 
Both fifth cranial nerves cut. Letters as in fig. 1. x ^. 



Experiment I. — Rabbit, decerebrate. Weight, 2300 g. Anaesthetic stopped, 10.45. 

11.14. 2 mg. tetra-methyl-ammonium chloride injected. Almost complete and im- 
mediate cessation of respiration for three seconds, followed by increased 
depth and frequency for ten seconds, and succeeded by a second almost 
complete paralysis lasting twenty-five seconds, then quick and good 
recovery. 

The blood-pressure (62 mm. Hg) showed a transient rise to 78 mm. Hg, 
followed by a moderately quick fall to 36 mm. Hg, with slowing of the 
frequency of the heart, and then a moderately quick rise to 1 1 1 mm. Hg. 
This rise, in turn, was followed by a gradual fall to 38 mm. Hg (at 11.18), 
and the fall by a gradual return to the normal height. 



28 PROFESSOR C. R. MARSHALL ON THE 

11.24. 2 mg. again injected. Respiration quickly depressed and paralysed for three 
seconds, followed by quick recovery and subsequently increased frequency 
and depth. 

The blood- pressure (70 mm. Hg) showed a very slight preliminary 
rise, followed by a fall to 39 mm. Hg and a secondary rise to 66 mm. Hg, 
and subsequent fall to 34 mm. Hg at 11.26. 

Experiment II. — Rabbit, decerebrate. Weight, 2100 g. Anaesthetic stopped, 11.30. 
Very low initial blood-pressure (26 mm. Hg). 

11.44. 1 mg. tetra-methyl-ammonium chloride into right external jugular vein. 
Slight gradual depression of respiration and then (thirty-five seconds 
after the injection) rapid paralysis. The blood- pressure gradually fell 
to 11 mm. Hg. Artificial respiration was performed for three minutes. 
The respiration was re-established, and the blood-pressure rose to 
104 mm. Hg. 

11.52^. 1 mg. again injected. Rapid and almost complete paralysis of respira- 
tion six seconds after commencement of injection with rapid recovery 
eleven seconds later. The blood-pressure fell from 106 mm. Hg to 
95 mm. Hg, and, after a transient rise to 102 mm. Hg, fell to 
91 mm. Hg. 

Experiment III. — Rabbit, decerebrate. Weight, 2025 g. Anaesthetic stopped, 3.5. 

3.31. 1 mg. tetra-methyl-ammonium chloride injected. Rapid, shallow, and irregular 
respirations almost immediately followed. Sixteen seconds later deep 
convulsive respirations occurred and were replaced, after thirty -five seconds' 
interval, by deep, regular respiration. 

The blood-pressure (80 mm. Hg) fell at first to 56 mm. Hg, and the 
heart-beats diminished to less than half their previous frequency. Five 
seconds before the commencement of deep breathing, the blood-pressure 
commenced to rise, and rose to 123 mm. Hg. With the onset of good 
regular breathing it gradually fell to 89 mm. Hg. 

3.37. 1 mg. again injected. Caused almost immediate paralysis of the respiration 
followed by gradual recovery. 

The blood-pressure fell from 84 mm. Hg to 58 mm. Hg, and 
the heart-beats from 13 to 5 in four seconds. The blood-pressure 
then rose to 103 mm. Hg and the heart-beats to 6^ in four seconds. 
At 3.39 the blood-pressure was 79 mm. Hg and the heart's frequency 9^, 
in four seconds. 

3.45. A third similar injection produced the same effects, but these were accompanied 
by marked muscular tremors. 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL- AMMONIUM COMPOUNDS. 29 

Experiment IV. — Eabbit, decerebrate. Weight, 1850 g. Fifth cranial nerves cut 
distally to Gasserian ganglia. Anaesthetic stopped. 2.54. Respiration irregular 
in rhythm. 

3.18. 0*5 mg. tetra-methyl-ammonium chloride injected. Transient stoppage of re- 
spiration followed by deeper, regular, and more rapid respiration. The 
blood-pressure fell from 80 mm. Hg to 68 mm. Hg, and with the onset 
of respiration rose to 108 mm. Hg. It then gradually fell to 72 mm. Hg 
(at 3.19£). 

3.20. 0*5 mg. again injected. Produced only slight depression of respiration. The 

blood-pressure was influenced as by previous injection, but to a somewhat 
less extent. 

3.21. 1 mg. injected. Produced paralysis of respiration for twenty seconds although 

blood-pressure was less influenced than by previous injections. 

Experiment V. — Rabbit, decerebrate. Weight, 2000 g. Both fifth cranial 
nerves cut. Anaesthetic stopped, 11.0. 

11.26. 2 mg. tetra-methyl-ammonium chloride injected. Respiration almost ceased 
for two seconds immediately after injection, then increased in depth and 
frequency for one minute, and subsequently returned gradually to normal. 

The blood-pressure (82 mm. Hg) rose slightly during the injection, 
and then, with marked slowing of the heart's frequency, fell rapidly to 
49 mm. Hg for a few seconds. A gradual rise to 71 mm. Hg followed, 
succeeded by a gradual fall to 5 1 mm. Hg and a gradual return to normal 
at 11.30. 

11.36. 1 mg. atropine injected. Slight transient stimulation of respiration and slight 

increase in frequency of heart. 

11.37. 2 mg. tetra-methyl-ammonium chloride injected. Respiration almost ceased 

for one second, then increased in frequency and gradually in depth for one 
minute, after which it gradually returned to normal. 

The blood-pressure showed successively a transient rise to 85 mm. Hg, 
a fall to 74 mm. Hg, a rise to 108 mm. Hg (at 11.38), a gradual fall to 
67 mm. Hg (at 11.39). 
11.40. l'5mg. tetra-methyl-ammonium chloride injected. Effects similar to previous 
injection, but somewhat greater diminution of frequency of heart. 

Experiment VI. — Rabbit, decerebrate. Weight, 2250 g. Both fifth cranial 
nerves cut. Anaesthetic stopped, 12.0. 

12.21. 1 mg. atropine injected slowly. 

12.25. 2 mg. tetra-methyl-ammonium chloride injected. Twelve seconds after com- 
mencement of injection respiration increased markedly in depth and some- 



30 PROFESSOR C. R. MARSHALL ON THE 

what in frequency. There was a simultaneous rise (occupying six seconds) 
of blood pressure from 114 mm. Hg to 150 mm. Hg, and after maintaining 
this level for twenty seconds it gradually fell. The effect on the respiration 
continued somewhat longer than that on the blood-pressure. The frequency 
of the heart was not affected. 

12.27. 2 mg. again injected. Respiration affected as after previous injection, but 
increase in depth somewhat greater. The blood-pressure fell from 124 mm. 
Hg to 102 mm. Hg for a few seconds, then rose to 156 mm. Hg, which 
was maintained for twelve seconds. It then fell, at first quickly, afterwards 
slowly, to 102 mm. Hg at 12.28|. 

12.29. 10 mg. tetra-methyl-ammonium chloride injected. Respiration completely 
paralysed seven and a half seconds after commencing the injection. No 
recovery occurred. 

Experiment VII. — Rabbit, decerebrate. Weight, 1850 g. Both fifth cranial nerves 
cut, the right not completely, proximal to the Gasserian ganglia. Anaesthetic 
stopped 10.50. 

11.21. 2 mg. tetra-methyl-ammonium chloride injected. Marked increase in depth of 
respiration commencing six seconds after beginning of injection and lasting 
four seconds and passing on to complete paralysis five seconds later. 
Spontaneous recovery did not occur, and artificial respiration only estab- 
lished infrequent respirations. 

The blood-pressure (96 mm. Hg) rose slightly and then fell rapidly to 
56 mm. Hg. Afterwards it rose to 98 mm. Hg and then fell to near the 
base line. 

Experiment VIII. — Rabbit, decerebrate. Weight, 1775 g. Left fifth 
cranial nerve cut. Anaesthetic stopped, 11.30. 

11.59. 1 mg. tetra-methyl-ammonium chloride injected. The respirations trebled in 
frequency a few seconds after the commencement of the injection, but 
rapidly diminished in depth and ceased ten seconds after beginning the 
injection. The paralysis lasted eleven seconds. After recommencing, the 
respiration quickly became efficient and remained somewhat more rapid 
although less deep for two minutes before gradually returning to normal. 

The blood-pressure (95 mm. Hg) showed a transient rise, then a quick 
fall to 76 mm. Hg, which was succeeded by a slight rise and then a gradual 
fall, concurrent with the recommencement of the breathing, to 56 mm. Hg. 
Recovery to the normal took several minutes. 

12. 4£. Injection repeated with similar result. 

12.29. 1 mg. again injected. A few very powerful respirations almost immediately 
followed, but the respirations quickly diminished in depth and ceased 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL-AMMONIUM COMPOUNDS. 31 

twelve seconds after the commencement of the injection. The paralysis 
continued for nine seconds, and then quickly became approximately normal 
for about six seconds, after which it became somewhat convulsive in type. 

The blood-pressure (104 mm. Hg) exhibited a transient rise, then a 
fall to 77 mm. Hg. With the recommencement of the respiration a slight 
transient rise occurred, followed by a fall to 39 mm. Hg, which was 
maintained. 

Experiment IX. — Rabbit, decerebrate. Weight, 1800 g. Both fifth cranial nerves 
cut, but left not completely. Anaesthetic stopped, 11.0. 

11.39. 1 mg. tetra-methyl-ammonium chloride injected. Respiration increased in fre- 
quency three seconds after the beginning of the injection, but diminished 
in extent, and four seconds later ceased. The paralysis lasted nine seconds, 
then respiration recommenced and rapidly became efficient, the depth 
being somewhat less than, but the frequency double, that before the injec- 
tion. The increased frequency continued for two minutes, then gradually 
passed away. 

The blood-pressure (73 mm. Hg) fell to 56 mm. Hg, and then com- 
menced to rise with the complete cessation of the respiration and continued 
until it reached 104 mm. Hg, just at the time when the respiration had 
become well re-established. It then fell gradually to 70 mm. Hg at 11.41. 

Experiment X. — Rabbit, decerebrate. Weight, 1900 g. Both fifth cranial nerves cut. 

Anaesthetic stopped, 10.40. 

11.4. 1 mg. tetra-methyl-ammonium chloride injected. Respiration diminished in 
extent five seconds after commencement of injection and was quickly 
followed by increasing frequency and gradually increasing depth for eight 
seconds. Then it diminished in extent and later increased until, forty- 
five seconds after the injection, it was 30 per cent, more rapid and the 
excursion of the lever was double that of the normal. Gradual return to 
normal then occurred. 

The blood-pressure (77 mm. Hg) showed a preliminary rise and then 
a prolonged fall, being 54 mm. Hg eleven seconds, and 40 mm. Hg two 
minutes, after the commencement of the injection. At three and a quarter 
minutes it was 49 mm. Hg, at six minutes 58 mm. Hg, and at thirteen 
minutes after the injection 77 mm. Hg. 

11.20. 2 mg. injected. Increased rapidity of respiration commenced four seconds 
after beginning of injection, but the curve gradually diminished in extent 
until twenty seconds later it was only one-seventh its previous height. 
The respiration then increased in extent, but diminished in frequency, 



32 PROFESSOR C. R. MARSHALL ON THE 

and twenty seconds later was almost normal. Afterwards it gradually 
became slower and finally ceased five minutes after the injection. 

The blood-pressure (77 mm. Hg) showed a slight preliminary rise and 
then a fall to 58 mm. Hg at ten seconds, and 49 mm. Hg at twenty 
seconds after the injection. No recovery occurred. 

Experiment XI — Rabbit, decerebrate. Weight, 2450 g. Both fifth cranial nerves 
cut proximal to Gasserian ganglia. Anaesthetic stopped, 11.20. 

11.36. 0'6 mg. tetra-methyl-ammonium chloride injected. Six seconds after com- 
mencement of injection the respiration increased suddenly to three times 
its previous frequency and about 20 per cent, in depth. This effect 
was maintained half a minute and then commenced to pass away, but the 
respiration had not reached its normal rate eight and a half minutes after 
the injection. 

11.44^. 1 mg. injected. The earlier part of the effects of this injection is shown 
in fig. 7. 

11.58. 1 mg. injected (took twelve seconds). Thirteen seconds after the commence- 
ment of the injection there occurred a marked increase in the frequency 
and a transient increase in the depth of the respirations, which soon 
commenced to pass away. 

11.594- 2 mg. injected (took eleven seconds) while respirations still rapid. Increased 
frequency and increased depth of respiration commenced three seconds after 
the beginning of the injection and was replaced by slowing and diminished 
depth five seconds later. After a further interval of five seconds the 
respiration ceased. The paralysis lasted for thirteen seconds. When the 
respiration recommenced, it was shallow and rapid, but it quickly reached its 
normal form. There was marked slowing of the heart and a fall of blood- 
pressure after the previous injection and a further slight slowing of the 
heart and slight fall of blood-pressure after this injection. 

12.5^. 5 mg. injected. The effect is shown in fig. 7. Occasional shallow respirations 
occurred afterwards. 

Experiment XII — Rabbit, decerebrate. Weight, 2200 g. Both Gasserian ganglia 

excised. Anaesthetic stopped, 3.38. 
3.53. 0*5 mg. tetra-methyl-ammonium chloride injected. Gradual diminution in depth 
of respiration commencing four seconds after beginning of injection and 
reaching maximal diminution, which was maintained six seconds, ten 
seconds later. Recovery gradually occurred during the next fifteen 
seconds. 

The blood-pressure (65 mm. Hg) rose slightly and then fell to 
45 mm. Hg nine seconds after the beginning of the injection, and to 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL-AMMONIUM COMPOUNDS. 33 

33 mm. Hg one minute after the injection. It showed practically no 
recovery. 
4.0^. 1 mg. injected. A rapid increase in depth and frequency occurred seven seconds 
after the beginning of the injection. Seven seconds later the frequency, 
and ten seconds later the depths of the respiration, began to diminish, and 
forty-five seconds from the commencement of the injection respiration 
ceased. After twenty seconds a few shallow respirations occurred at 
intervals during the next half-minute. The blood-pressure very gradually 
fell to 15 mm. Hg. 
Cause of the Respiratory Paralysis. — As previously stated, Iodlbauer # ascribes 
the temporary paralysis of the respiration to stimulation of the terminations of the 
fifth cranial nerves in the nose. He came to this conclusion because (l) he observed 
closure of the glottis— an associated act of the Kratschmer-Hering reflex — during the 
respiratory standstill ; (2) he was unable to produce respiratory paralysis after anaesthetis- 
ing the nasal mucous membrane ; (3) he did not observe the characteristic effect in 
cats, animals in which the typical Kratschmer-Hering reflex is absent. In these 
animals he obtained transient powerful stimulation of the respiration followed by some 
depression and rapid return to the normal. 

It is difficult to understand why Iodlbauer failed to obtain the respiratory 
paralysis in cats. It is possible that the animal used for the experiment in which 
efficient doses were given may have been relatively insusceptible to tetra-methyl- 
ammonium chloride, since variations in the reaction to this substance occur in these 
animals and also in rabbits. In the few experiments I have made on cats, respiratory 
paralysis has been as easily induced as in rabbits, although generally somewhat larger 
relative doses were necessary for the purpose (see fig. l). 

Nor have I been able to corroborate Iodlbauer's statement that anaesthetising 
the nasal mucous membrane of rabbits prevents the appearance of the respiratory 
paralysis. Even after the careful injection of large quantities of a 5 per cent, solution 
of cocaine hydrochloride so as to reach the whole mucous membrane, the intravenous 
injection of 1 mg. tetra-methyl-ammonium chloride produced the same effect as before 
the application of the cocaine. 

It would seem further that if this action is due to stimulation of the trigeminal 
nerve-endings in the nose, the local application of such a diffusible substance as tetra- 
methyl-ammonium chloride would induce it. As I shall show in a later communication, 
the application of 1 in 5000 solutions of this substance to frogs' muscles produces a 
maximal effect within a few seconds, and it is inconceivable that the nasal mucous 
membrane should act as a semi-permeable membrane to it. Yet the injection of 1 to 2 
per cent, solutions into the nasal cavities of anaesthetised animals, the respiration of 
which was paralysed by intravenous injections of 1 mg., produced no effect whatever. 
Iodlbauer apparently used this method with equivocal results, which he attributes to 

* hoc. cit. 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 2). • 5 



34 



PROFESSOR C. R. MARSHALL ON THE 



the difficulty of avoiding mechanical stimulation of the mucous membrane by the 
instillation. 

Moreover, as is evident from the experiments already summarised (Exps. IV., V., 
VI., X., XL, XII.), the effect is still obtained after section of both fifth cranial nerves 
in the base of the skull. And in one experiment in which these nerves were divided in 
the interval between two injections the effect produced by the injections was similar 
in both cases (fig. 8). 

It would therefore seem that the cause of the temporary paralysis of the respiration 
is not to be found in stimulation of the nerve-endings in the nose. As previously 
stated, it is in all probability due mainly to transient depression or paralysis of the 
myo-neural junctions in the respiratory muscles. It has already been shown that 




Fig. 8.— Effect of tetia-niethyl-anunoiiium chloride before and after division of both fifth cranial nerves in decerebrate 
rabbit. 12.12, right fifth nerve cut ; 12.13 left nerve cut. Letters as in previous figures, x |. 

greater or less paresis of the motor nerve-endings to the sterno-mastoid and the muscles 
of the fore limbs occurs a few seconds after the injection of a dose of tetra-methyl- 
ammonium chloride sufficient to paralyse the respiration. An examination of the 
various experiments in which the effect on the respiration and the excitability of 
the fore-limb muscles was observed shows that the paresis of the fore-limb muscles 
is synchronous with the paralysis of the respiration. This is seen in fig. 1 in the 
cessation of the spontaneous muscular movements, and also in fig. 2, although the 
respiration in the latter case was not so quickly paralysed. It was further found 
that the duration of the paralysis in each case was similar. After some injections 
the limb muscles seemed to be more influenced than the respiration — a weak stimulus 
which had previously produced a good contraction inducing for a short time no reaction 
after a dose sufficient to depress, but insufficient to paralyse, the respiration : or, as 
shown in fig. 1, failing to react to the same degree as before, after the respiration 



PHARMACOLOGICAL ACTION OP TETR A- ALKYL- AMMONIUM COMPOUNDS. 35 

had completely recovered from the paralysis ; but even in these cases the depression or 
paralysis of the motor nerve-endings and of the respiration was shown to run a parallel 
course. When the influence of tetra-methyl-ammonium chloride on the terminations 
of the phrenic nerves was investigated, complete agreement was found to exist between 
their irritability and the effect on the respiration. In these experiments the phrenic 
nerves were isolated in the base of the neck and one or other, usually the right, 
stimulated in continuity. The respiration was recorded by means of a phrenograph 
connected to a tambour recorder. A considerable number of such experiments were 
made, but as they all conform to that shown in fig. 1, it is only necessary to refer to 
this. In this experiment the right musculo-cutaneous and right phrenic nerves were 
stimulated simultaneously. As will be seen from the tracing (line R), the effect on 
the respiration and the effect of stimulating the phrenic are completely concordant. 

It would seem therefore that the respiratory paralysis produced by tetra-methyl- 
ammonium chloride is largely, if not solely, due to paralysis of the nerve endings in 
the respiratory muscles. This being the case, we might expect a comparable effect to 
be given by curare. This substance, however, is somewhat slower in its action and 
much more prolonged in its effects than tetra-methyl-ammonium chloride, and after 
the injection of doses sufficient to paralyse the respiration spontaneous recommencement 
of the breathing cannot be obtained without artificial respiration. According to Tillie,* 
the smallest dose of curarin, administered intravenously, which will completely paralyse 
a rabbit is 0*2 mg. p. kg. body-weight; 0*3 mg. to 0*5 mg. p. kg. cause loss of reflexes 
and spontaneous breathing for ten to fifteen minutes. Consequently in an experiment 
on a cat — animals which, according to BoEHM,t are somewhat less sensitive to curarin 
than rabbits — carried out in a similar manner to those referred to on p. 18, anaesthesia 
being maintained throughout the experiment, a dose of 0"16 mg. p. kg. body-weight 
was decided on. The intravenous injection of this quantity of curarin caused gradual 
failure of the respiration commencing twenty-one seconds after the beginning of the 
injection and going on to complete paralysis two minutes later. Electrical stimulation 
of one of the cords of the brachial plexus and of the right phrenic nerve showed 
gradually diminishing contraction of the fore limb and the diaphragm respectively 
pari passu with the failure of the respiration. With complete cessation of the 
respiration stimulation of the phrenic nerve caused no contraction of the diaphragm, 
but stimulation of the brachial cord produced a just appreciable movement of 
the limb4 Artificial respiration was performed, and five minutes later spontaneous 
breathing recommenced, but even with the aid of short periods of artificial respira- 

* Arch.f. exp. Path. u. Pharm., xxvii. p. 22 [1890]. 

t Festschrift zu G. Ludwig's 70. Geburstag, 1886, quoted by Tillie, loc. cil., p. 4. 

% This result although slight is probably unusual. By careful dosage of curarin Tillie was able to keep an 
animal paralytic without stopping the respiration ; and A. Mosso (Memoria della R. Acad, delle Scievza di Torino, ser. 
ii. vol. liii. p. 397, 1903) and M. Chio (Arch, di Farmacol. speriment. e Scienz. affin., xii. p. 533, 1912), by comparing 
the respiratory movements of the thorax and abdomen, have shown that the muscles of the thorax are paralysed 
before those of the diaphragm. 



36 PROFESSOR C. R. MARSHALL ON THE 

tion nearly twenty minutes elapsed after the paralysis of the respiration before 
practically normal respiration was established, and it was some minutes later 
before the diaphragm reacted to stimulation of the phrenic to the extent it did 
previous to the injection. After the injection of double this dose, paralysis of the 
muscles and respiration appears much more quickly, but three to four times the dose 
mentioned is necessary to produce that rapid paralysis of the respiration which is seen 
after tetra-methyl-ammonium chloride. Owing to the prolonged action of curarin on 
the myo-neural junctions, a strict comparison with the paralysing action of tetra-methyl- 
ammonium chloride on the respiration is not possible ; but it is interesting to note that 
the type of the paralysis is the same in both cases and that after large doses of the 
two substances the paralysing effect on the respiration is practically identical. In view 
of the convulsant action of tetra-methyl-ammonium chloride, it is further of interest to 
note that Tillie # observed weak convulsions, apparently on non-anaesthetised animals, 
after the injection of 1 mg. to 2 mg. curarin prior to the appearance of complete 
paralysis, and these were more marked when artificial respiration was performed 
throughout the experiment. 

There still remains the question whether the cause of the respiratory paralysis is 
wholly due to the paresis of the nerve endings in the respiratory muscles. As previously 
mentioned, PoHL,t working with some quaternary papaverine derivatives, observed 
this form of temporary expiratory paralysis with spontaneous recovery after intravenous 
injections into rabbits. And yet he found that these substances had no paralysing 
action on the motor nerve-endings of frogs. He came to the conclusion that the effect 
on the respiration was due to an action on the respiratory centre, and he explained the 
effect of tetra-methyl-ammonium chloride in the same way. But the only experiment 
he appears to have made was one to controvert the view of Tappeiner, and in this he 
divided the ophthalmic branches of the fifth cranial nerves, and still found that he 
obtained the characteristic paralysis of the respiration after the intravenous injection of 
this substance. Its action was not further investigated. 

In order to determine whether the paralysing action on the respiration produced by 
tetra-methyl-ammonium chloride is partly central, two experiments in which the injec- 
tions were made alternately into the carotid artery and the facial vein were performed 
on anaesthetised rabbits. Although the concentration of tetra-methyl-ammonium 
chloride reaching the respiratory centre after injection into the artery must have been 
considerably greater than after injection into the vein, a larger dose was necessary to 
induce respiratory paralysis. Thus the injection of 1 mg. into the carotid artery pro- 
duced in both animals merely slight gradual diminution of depth and slowing of the 
respiration, whereas this dose, when injected into the facial vein, caused almost complete 
temporary paralysis. When double the dose injected into the vein was injected into 
the artery, almost identical effects on the respiration were obtained from the two 
injections. Some difference was noted in the initial effects on the blood-pressure, the 

* hoc, cit., ]>. 22. t Loc. cit. 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL- AMMONIUM COMPOUNDS. 37 

fall being more gradual and the slowing of the heart's frequency much less marked 
after the intra-arterial than after the intravenous injection. 

The paralysing action of this substance on the respiration would therefore seem not 
to be due to an action on the respiratory centre. Nor is it due to an action on the 
circulation or on the peripheral nervous mechanism within the lungs, since, as will be 
shown later, it is much more transient than the circulatory action, and it is obtained 
after section of both vagi or after the injection of sufficient atropine to paralyse the 
vagal terminations. 

That tetra-methyl-ammonium chloride is not without action on the respiratory centre 
is evident from the experiments on decerebrate animals. But instead of being 
depressant to this centre, it is powerfully stimulant. And that the paralysis is not 
due to exhaustion following stimulation is obvious from the fact that stimulation of 
the respiration is usually present in decerebrate animals after the paralytic stage has 
passed away. The larger doses of tetra-methyl-ammonium chloride which are some- 
times necessary to paralyse the respiration of decerebrate animals, and the more rapid 
recovery from the paralysis, also point to this conclusion. It has been mentioned that 
strong or moderately strong electrical stimuli applied to nerves may still produce 
contraction of the innervated muscles after doses of tetra-methyl-ammonium chloride 
sufficient to paralyse the respiration, although moderately weak stimuli may no longer 
do so, and there can be little doubt that a powerfully stimulated respiratory centre acts 
in a similar manner, only more efficiently. In other words, the impulses from a 
respiratory centre in this condition are able to overcome a greater degree of depression 
in the peripheral structures, and hence the depression must be made more profound by 
the injection of larger doses in order to obtain paralysis ; and as the stimulation continues 
for a much longer time than the paralysis, the more rapid recovery may be similarly 
explained. It is probable that the convulsant action noticed after toxic doses to normal 
animals has a similar explanation. That the stimulation of the respiratory centre does 
not occur in anaesthetised animals when the substance is administered intravenously is 
due to the well-known fact that it is more difficult to stimulate this centre in anaesthetised 
than in normal animals, and any slight stimulation which may be induced would, in 
this case, be checked by the simultaneous paralysing action on the myo-neural junctions. 
By the injection of small doses into the carotid artery of anaesthetised animals, slight 
stimulation of the respiration, both increase in depth and in frequency, can be 
demonstrated. 

Influence of the Vagus on the Respiratory Paralysis. — Anaesthetised animals in 
which both vagi have been divided are more sensitive to the action of tetra-methyl- 
ammonium chloride than animals with the vagi intact. Thus a dose which may 
produce in an ordinary anaesthetised animal slight temporary depression without 
cessation of the respiration will, after section of both vagi, cause complete temporary 
paralysis ; or a dose sufficient to paralyse for a few seconds the respiration of an animal 
with undivided vagi, may, after the vagi have been cut, produce permanent respiratory 



38 PROFESSOR C. R. MARSHALL ON THE 

paralysis. If not, the period of paralysis is longer than before the vagi were severed 
and the period of recovery of the respiration is prolonged. Decerebrate animals in 
which the section was made above the anterior quadrate bodies also proved more 
sensitive to the paralysing action of tetra-methyl-ammonium chloride after the vagi 
were cut than before. When the section through the mid-brain was made behind these 
bodies, division of the vagi was followed by irregular, spasmodic, and inefficient 
breathing, which gradually failed. All that could be deduced from the injections of 
tetra-methyl-ammonium chloride in these vagotomised animals was that they produced 
no stimulation of the respiration. This apparently increased sensitiveness of vago- 
tomised animals to the influence of tetra-methyl-ammonium chloride is doubtless due 
to the fact that after division of the vagi the centripetal impulses produced by 
inspiration and expiration, which normally play an important part in the regulation of 
the respiration, are cut off, and the respiratory centre is thus deprived of one of its 
most efficient forms of stimulation to activity. After doses of atropine sufficient to 
paralyse the pulmonary vagus, no increased sensitiveness to tetra-methyl-ammonium 
chloride was observed, which is probably due, as some of my experiments seem to 
suggest, to a limited antagonism existing between these substances. 

The experiments which have been described prove that the action of tetra-methyl- 
ammonium chloride in paralysing the respiration is fundamentally due to a depressant 
action on the myo-neural junctions of the respiratory muscles. It will be shown in 
subsequent communications that the paralysing effect on the respiration of other tetra- 
alkyl-ammonium compounds is proportional to their power of causing muscular paresis. 

Effect on the Circulation. 

In anaesthetised animals the intravenous injection of tetra-methyl-ammonium 
chloride invariably causes a fall of blood-pressure and almost invariably a decided 
diminution in the frequency of the heart-beats (fig. 6). The fall of blood-pressure, 
occasionally after an insignificant rise, begins a few seconds after the commencement of 
the injection and usually about two seconds before the respiration is influenced. At 
first it is generally rapid, and later more gradual. After a dose of 1 mg. the fall may 
occupy twenty seconds and the blood-pressure may then be less than half its original 
height. If the respiration is inefficient, a slight asphyxial rise may occur, but commonly 
this is absent and the low blood-pressure continues, notwithstanding an efficient 
respiration, for one to three minutes and then gradually returns to normal. In a few 
cases after the administration of this dose the blood-pressure has fallen to a low level 
and no recovery has occurred. 

The fall in the frequency of the heart-beats runs a similar course, but is of 
somewhat shorter duration. It commences with the fall of blood-pressure and may 
amount to '25 per cent., but is generally less than this. It begins to pass away before 
the blood-pressure commences to rise, and has usually completely passed away before 
the blood-pressure has reached its normal height. It is apparently wholly due to 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL-AMMONIUM COMPOUNDS. 39 

stimulation of the vagal endings in the heart, since it is unaffected by division of the 
vagi and is abolished by the injection of sufficient atropine to paralyse these endings. 
After paralysing the vagal endings by means of atropine, the injection of 1 mg. tetra- 
methyl-ammonium chloride still causes a fall of blood-pressure, but the fall is more 
gradual and is usually much less in extent that in non-atropinised animals. Otherwise 
it runs a similar course. In cats after minimal doses the fall of blood-pressure and of 
the frequency of the heart is usually less than in rabbits, but otherwise the effects 
produced are the same. 

After larger doses the fall of blood-pressure is more rapid, more marked, and more 
prolonged, and unless the respiration becomes efficient or artificial respiration is per- 
formed, no recovery follows. In some cases even when spontaneous breathing has 
recommenced or artificial respiration has been performed, the blood-pressure showed no 
recovery or rose but slightly. In these the blood-pressure remained at a very low 
level and the animal eventually succumbed. 

In decerebrate animals a fall of blood-pressure with slowing of the heart also 
occurs (figs. 7 and 8), but is almost invariably preceded by a slight transient rise, also 
associated with a diminution of the heart's frequency. With the fall of blood-pressure 
the heart for a short period often becomes very slow. The vagal endings are also more 
sensitive to stimulation than normally — a stimulus which before the injection produced 
no effect causing after the injection a marked vagal action. The fall of blood-pressure 
is succeeded by a rise, often considerably above the normal height, which is soon 
followed by a fall below normal, the rise and fall occupying about twenty seconds. 
After a varying interval gradual recovery occurs as in anassthetised animals. When 
atropine has been given previously, there is usually little or no preliminary fall of 
blood-pressure after the injection of small doses of tetra-methyl-ammonium chloride, 
and after a short interval a considerable rise occurs, which is maintained for about 
twenty seconds, and is followed by a fall to normal or a little below normal (fig. 5). 
Iodlbauer # also observed a rise of blood-pressure in non-anaesthetised animals after 
atropine had been given, but this was only followed by a fall below normal when large 
doses of tetra-methyl-ammonium chloride were injected. 

The effect of tetra-methyl-ammonium chloride on the circulation is somewhat 
complex. Iodlbauer, as a consequence of the experiment just mentioned, concludes 
that it causes a marked stimulation of the vaso-motor centre, and, since he obtained 
less slowing of the heart's frequency after than before division of the vagi, that it also 
stimulates the cardio-inhibitory centre. The variable effects he obtained on the blood- 
pressure he explains by stating that sometimes stimulation of the vagal mechanism, 
sometimes stimulation of the vaso-motor centres, is predominant. The fall of blood- 
pressure, apart from that caused by stimulation of the vagus, he believes is due to 
paralysis of the vaso-motor centres. In my own experiments the stimulation of the 
cardio-inhibitory centres was relatively slight ; after division of the vagi in decerebrate 

* hoc. cit., p. 196. 



40 PHARMACOLOGICAL ACTION OF TETRA-ALKYL-AMMONIUM COMPOUNDS. 

animals the slowing of the frequency of the heart was only 10 per cent, less than 
before the vagi were divided. The greater vagal effect seen in decerebrate animals as 
compared with anaesthetised animals is mainly due to greater stimulation of the vagal 
terminations. 

The fall of blood- pressure in anaesthetised animals is mainly, and the early fall of 
blood-pressure in decerebrate animals is wholly, vagal in origin, and is due to stimula- 
tion of the vagus endings in the heart ; but the late fall in decerebrate animals and the 
fall in anaesthetised animals are partly vascular, and appear, contrary to Iodlbauer's 
statement, to be peripheral, and not central, in nature. This was shown by observing 
the effects of stimulating a splanchic nerve on the blood-pressure. Thus in a small 
etherised rabbit (weight 1600 g.) stimulation of the left splanchnic nerve raised the 
blood-pressure from 52 mm. Hg to 72 mm. Hg. After the injection of 1 mg. tetra- 
methyl-ammonium chloride stimulation of the nerve with the same stimulus produced 
no effect until the blood-pressure commenced to rise. Two minutes after the injection 
it raised the blood-pressure from 35 mm. Hg to 37 mm. Hg ; four minutes after the 
injection from 44 mm. Hg to 50 mm. Hg ; and six and a half minutes after the injection 
from 52 mm. Hg to 65 mm. Hg. Whether the effect is due, as Tillie # maintained for 
curarin, to a depressant influence on the vasoconstrictor nerve endings or upon the 
muscle of the vessel walls, has not been determined. The perfusion of the blood-vessels 
of winter frogs has produced, with concentrations down to 1 in 400,000, only contrac- 
tion, whereas plethysmographic experiments with the intestines of etherised rabbits 
have shown only dilatation of the vessels. It seems probable that the dilatation of the 
blood-vessels is one affecting mainly the splanchnic area and is due to a depressant 
influence on the terminations of the splanchnic nerves. This is probably the main 
vascular factor contributing to the fall of blood-pressure, but the diminished resistance 
associated with the paralysed condition of the voluntary muscles may not be without 
influence. 

The temporary rise in blood-pressure occurring in atropinised decerebrate animals 
and the rise following the preliminary fall in non-atropinised decerebrate animals are 
due to stimulation of the vaso-motor centres. The curve of the rise and fall in non- 
atropinised animals strongly suggests an asphyxial origin, but while there can be no 
doubt that the increased carbon dioxide tension in the blood caused by the paralysis 
of the respiration acting upon an excited vaso-motor centre is partly responsible for 
the rise, it cannot be wholly due to this cause, since it occurred in similar form in 
animals with an efficient respiration (see fig. 7, in which, however, the rise is relatively 
small). Moreover, it only followed the earlier injections of an experiment, and was 
less marked or absent after moderate or large doses had been given. The rise of blood- 
pressure may therefore be attributed to stimulation of the vaso-motor centres and the 
subsequent fall to the increasing preponderance of the peripheral action already 
referred to. 

* Loc. cit., p. 29. 



( 41 ) 



III. — Polychaeta of the family Nereidse, collected by the Scottish National 
Antarctic Expedition (1902-1904). By L. N. G. Ramsay, M.A., B.Sc, Carnegie 
Research Scholar, Christ's College, Cambridge. Communicated by Dr. J. H. 

ASHWORTH. 

(MS. received October 7, 1913. Read December 15, 1913. Issued separately March 30, 1914.) 

[Plate III.] 

The collection of Nereidse brought home by the Scotia proves to be of considerable 
interest. As other expeditions have indicated, the family is but poorly represented in 
the antarctic or sub-antarctic regions ; and although a large number of specimens were 
collected at the South Orkney Islands, these have all proved to belong to one species, 
N. kerguelensis M'Int. No nereids were obtained at any of the deep-water stations 
farther south — the family being decidedly littoral in its range. 

The chief interest, however, lies in the material collected so assiduously throughout 
the vessel's wanderings. Six other species were obtained, including one from the 
Falkland Islands, hitherto und escribed. 

The investigations of the more recent workers on the Nereides continually tend to 
widen the distribution of hitherto known species, and to link up forms which have been 
described as distinct — in some cases apparently from the supposition that it was incon- 
ceivable that one and the same species could inhabit areas so widely separated as, say, 
the Indian Ocean and the North Atlantic. It is becoming more and more evident, 
however, that in the determination of species, locality must not be taken into account. 
Several species are already recognised as having a range that is practically cosmopolitan 
(e.g. N. dumerilii, from the North Atlantic, North Pacific, and Persian Gulf; N. mirabilis, 
from the West Indies, Red Sea, Persian Gulf, and Malay Archipelago). It is therefore 
clear that the world-wide distribution of any particular species is a contingency that 
must be reckoned with. 

The means of dispersal which render this world-wide distribution possible can only, 
for the present, be a matter of surmise ; the carriage of pelagic larvse by ocean currents 
may be sufficient to account for it ; and in this connection, also, the investigations of the 
Scotia naturalists supply a very suggestive hint in the discovery of nereids living and 
breeding among floating gulf-weed in mid-Atlantic. These, it may be noted, have been 
described as a variety of the species N. dumerilii. 

The excellent condition of the Scotia material has rendered its examination and the 

determination of the species comparatively easy. The descriptions following are in no 

case based on a single specimen ; where only a few specimens were found, these were all 

examined in detail ; where examples were numerous, the individual variation has been 

taken into account. This is of considerable importance, as the great latitude of indi- 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 3). 6 



42 



MR L. N. G. RAMSAY ON POLYCHiETA OF THE FAMILY NEREIDS, 



vidual variation, in such characters as the numbers and grouping of paragnaths, and the 
relative lengths of appendages, has apparently led astray several former investigators of 
the group, and has resulted in the creation of a deplorable number of perfectly indistin- 
guishable species. This is undoubtedly due in some measure to the laborious nature of 
the examination necessary for the adequate determination of the characters of a nereid. 

In the preparation of this report, the extensive collections of Nereidse in the 
Cambridge University zoological laboratories, hailing from all regions of the globe, have 
been of great assistance ; for these I am indebted to Mr F. A. Potts. It may be 
mentioned here that the writer is at present engaged on a revision of the family as a 
whole, and on this account one or two points, such as the relationships of the species 
allied to N. falklandica, or of the species comprised in the sub-genus or group Platy- 
nereis, have not been discussed in the present paper. 

I have to thank Professor J. Stanley Gardiner and Mr F. A. Potts for helpful 
advice. My best thanks are also due to Professor F. Jeffrey Bell for kindly allowing 
me access to the collections at the Natural History Museum for purposes of comparison ; 
finally, to Dr W. S. Bruce himself for permitting me to examine the material. 



Table of Species, with Localities. 



Nereis herguelensis M'Int. 

Nereis eugenise (Kbg.). . 
Nereis mirabilis Kbg. 
Nereis pelagica Lin. 
Nereis falklandica n. sp. 
Nereis australis (Schmarda). 



Nereis dumerilii Aud. et Edw.. 
nov. comata. . 



subsp. 



South Orkneys. 
Falkland Islands. 
Falkland Islands. 
Brazilian coast. 
Cape of Good Hope. 
Falkland Islands. 
Falkland Islands. 
Cape of Good Hope. 

North Atlantic (mid-ocean). 



Nereis herguelensis M'Int. 
Nereis herguelensis M'lntosh (10), p. 225, pi. xxxv. figs. 10-12. 

Station 325. Scotia Bay, South Orkneys ; many specimens. 

Station 118. One small specimen, Port Stanley, Falkland Islands, January 1903. 

This species was the only representative of the Nereidse obtained at the South 
Orkneys, where it was apparently fairly common in the littoral zone at Scotia Bay. 
Many specimens were collected in 9-10 fathoms; they are accompanied by tough 
membranous tubes in which they lived. 

Two specimens from Scotia Bay (season ?) are undergoing change to the heteronereid 
form. 

It appears to be a peculiarity of this species that the spinigerous setae of the lower 



COLLECTED BY THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 43 

neuropodial bundle are homogomph in form. This, at any rate, is constant in the 
Falkland Islands specimens, as well as in those from the South Orknejs. 

N. kerguelensis has a wide distribution in the sub-antarctic regions, having 
been recorded from Kerguelen, South Georgia, the Falkland Islands, and the South 
Shetlands and Graham Land, besides the South Orkneys. Kecently Von Marenzeller 
(12, p. 15) has recorded specimens from the eastern Mediterranean, collected by the 
Austrian oceanographic vessel Pola, and he mentions also some from the Azores in 
the collections of the Prince of Monaco. 

Nereis eugenise (Kbg.) Char, emend. Ehl. 

Nereis eugenise, Ehlers, (3), p. 67, pi. iv. figs. 94-105. 

Station 349. Two specimens, dredged in 15 fathoms, sandy bottom, off Tussock 
Islands, Falkland Islands, December 2, 1903. 

These are both large specimens, and agree in all respects with Ehlers' full descrip- 
tion and figures. 

This species has been recorded from the shores of various parts of the southern 
extremity of South America, but not previously from the Falkland Islands. 

Nereis (Ceratonereis) mirabilis Kbg. 
Nereis {Ceratonereis) mirabilis, Ehlers, (2), p. 117, pi. xxxvii. figs. 1-6. 

Station 81. One specimen, dredged in 36 fathoms, coral bottom, about 200 miles 
off the coast of Brazil, 18° 24' S., 37° 58' W., December 20, 1902. 

It is a small example, incomplete posteriorly, and showing the characters typical 
of the species. 

The type specimen of N. mirabilis was obtained off the Brazilian coast in 9° S. 
latitude (9, p. 170). The species has subsequently been recorded from Florida 
(2, p. 117), the Red Sea (7, p. 172), Porto Rico and Bermuda (13, p. 193), Amboina 
(14, p. 336), and the Persian Gulf (15, p. 392). 

Nereis pelagica Lin. (PL III. figs. 1, 2.) 
Nereis pelagica M'Intosh (11), p. 267, and figs. 
Station 478. One example from the shore, Table Bay, South Africa, May 14, 1904. 
Station 483. Two examples, trawled in 25 fathoms, sand and kelp, entrance to 

Saldanha Bay, South Africa, May 21, 1904. 
The three are all small specimens, the largest being 25 mm. long; fortunately 
the proboscis is in all cases everted, and this greatly facilitates examination. 

I can find no differences of importance between these and British specimens of 
N. pelagica ; the general form of the body, the parapods, setse, and the prostomium 
and its appendages all agree, as does the armature of the proboscis, except that the 



44 MR L. N. G. RAMSAY ON POLYCHJETA OF THE FAMILY NEREIDS, 

ventral paragnaths of the basal ring ( VII.-VIII.) do not exhibit an anterior row of 
markedly larger points, such as is usual in examples from Britain (v. figs. 1 and 2). 

The noto-cirri are rather shorter than in typical British specimens, but nevertheless 
overreach the noto-ligules all along the body. 

N. lucipeta Ehl., from Great Fish Bay, some distance farther north, very 
closely resembles the present species, as Ehlers himself states (6, p. 71). Ehlers' 
specimens were all in the heteronereis condition, and his main grounds for separating 
the species from pelagica were : the swollen outline of the noto-cirri of the first seven 
pairs of parapods, the number of parapods in the anterior (unaltered) portion of the 
body, i.e. 19-20 pairs, and the longer tentacular cirri (the longest reaching to the 
9th— 1 1th setigerous segment). 

I have before me a small example of A/, pelagica, obtained at Plymouth in March 
1913, 25 mm. in length, and in the heteronereis condition ; this specimen, as regards 
the parapods and their modifications, exactly agrees with Ehlers' description and 
figures of N. lucipeta. The change in the form of the parapods occurs at the 21st pair, 
and the lst-7th pairs — of the anterior region — have the basal part of the noto-cirrus 
swollen to a sausage shape, and terminate in a short, unswollen tip, exactly as figured 
by Ehlers for N. lucipeta. 

The length of the tentacular cirri is not of much importance, as it is subject to 
considerable variation. 

While the present record forms an important extension of the known range of 
N. pelagica, it is not surprising that the species should be found to occur on the 
coasts of South Africa, as it has previously been reported from such widely separated 
regions as northern Europe, the northern Pacific (both the Japanese and American 
shores), and the west coast of South America. 

Nereis (Perineris) falklandica, n. sp. (PI. III. figs. 3-10.) 

Station 118. One specimen, Port Stanley, Falkland Islands. 

Station 349. One specimen, dredged off Tussock Islands, Falkland Islands, in 
15 fathoms, sandy bottom, December 2, 1903. 

These two are in excellent preservation, but one is incomplete posteriorly. The 
complete example measures 80 mm. in length by 6 mm. in width over parapods. 
There are 93 setigerous segments ; the width hardly decreases till the 80th, behind 
which the body rapidly tapers. The other example is smaller. 

The anal cirri are very short, as are also the peristomial cirri. In the preserved 
animal (in alcohol) the tentacles and peristomial cirri are livid, in marked contrast to 
the deeply-pigmented prostomium ; the palps and the basal divisions of the peristomial 
cirri are intermediate in colour, or slightly pigmented. The eyes are unusually small, 
and each is surrounded by a pale ring. In the incomplete specimen the pigment is 
more developed than in the larger one, and the dorsal surface of the body-segments 



COLLECTED BY THE SCOTTISH NATIONAL ANTAKCT1C EXPEDITION. 45 

is dull purplish-brown like the head. In the other the body is practically unpigmented, 
although the head agrees with the above description (v. fig. 3). 

The proboscis (figs. 4, 5) was in both cases retracted, but owing to the good 

preservation of the specimens, examination by dissection gave very satisfactory results. 

The two were found to be identical as to the form and arrangement of the paragnaths. 

These, numerous, black and conical, and of very diverse size, are arranged as follows : — 

I. An irregular area of numerous very minute black points, with a few larger 

ones situated medio -anteriorly. 

II. A closely-packed triangular group of about fifteen large paragnaths. 

III. A small close-set transverse group of about ten, of fairly large size. 

IV. About twenty-five, of varying size, forming a transversely-elongate group. 
V. One large median paragnath. 

VI. One large paragnath, slightly elongate transversely. 

VI I. -VIII. A wide band of very numerous cones, smallest and most numerous in 

the mid- ventral region, becoming fewer and larger towards the dorsum and 

also towards the anterior margin of the belt, and extending dorsalwards 

nearly to VI. 

The jaws are nearly black, opaque, with one big " tooth " or notch near the tip, the 

remainder of the cutting edge being only slightly crenate (fig. 6). 

In the anterior region of the body the lobes of the parapods are short, stout, 
rounded, and of more or less even size. Proceeding towards the posterior end, a 
gradual transition of form takes place ; the base of the dorsal ligule becomes enlarged 
and more enlarged, assuming an oblong form and carrying the cirrus with it ; the 
rounded tip of the ligule projecting below the cirrus becomes less prominent. At the 
same time the lower lobes of the parapod are reduced in size, especially the ventral 
ligule (figs. 8-9). Thus the outline of the parapods changes gradually from a quad- 
rangular to a triangular one. 

The setse, numerous, short, closely-packed, and of a horny yellow colour, are thus 
distributed :■ — 

A. Notopodial bundle : homogomph spinigers. 

B. Upper neuropodial bundle : homogomph spinigers (above). 

,, ,, heterogomph falcigers (below). 

C. Lower neuropodial bundle : heterogomph falcigers. 

The heterogomph spinigers which appear on the upper margin of C in most species 
of Nereis have apparently been eliminated. The bristles do not vary appreciably in 
form from the 1st parapod to the 70th at least. The falcigers have stout shafts, and 
short stout appendages (fig. 10). 

In the general external form, and in the characters of the parapodia, setae, and the 
prostomium and its appendages, the present species closely resembles N. variegata Gru., 
and others of the same group : N. marionii A. et E. (11, p. 295), N. melanocephala 
Mint. (10, p. 216), N. (Pseudonereis) anomala Grav. (7), N. ruficeps Ehl. (6), 



46 MR L. N. G. RAMSAY ON POLYCH^ETA OF THE FAMILY NEREIDS, 

N. gaUapagensis Kbg. (8). From all these, however, the armature of the proboscis 
exhibits considerable differences. No other species of nereid, to my knowledge, exhibits 
a similar arrangement in group I., i.e. very numerous, minute, scattered conical parag- 
naths. Again, there is in N. falhlandica none of the markedly pectiniform arrange- 
ment of the paragnaths of groups II., III., and IV. exhibited by N. variegata, 
N. anomala, and N. gaUapagensis. VII.-VIII. also are distinctive. 

This group of species, in common with the rest of the family, stands in considerable 
need of revision. In the case of N. variegata, two different types appear to have been 
confused under the one name by different authors. 

Nereis {Platynereis) australis (Schmarda). 
Nereis {Platynereis) australis, Ehlers (5), p. 26, pi. iii. f. 16-20, iv. f. 1-2. 
Station 349. One specimen, dredged in 15 fathoms, sandy bottom, off Tussock 

Islands, Falkland Islands, December 2, 1903. 
Station 118. One specimen, Port Stanley, Falkland Islands, January 1903. 
Station 478. One specimen, shore, Table Bay, South Africa, May 1903. 
Station 482. Four examples, in 2-8 fathoms, shells and sand, Houtjes Bay, 

Saldanha Bay, South Africa, May 21, 1903. 
Station 483. Numerous examples, in 25 fathoms, sand and kelp. Entrance to 

Saldanha Bay, South Africa, May 21, 1904. 
The Scotia specimens are all small, none exceeding 4 mm. in width over pararodia, 
by about 50 mm. in length. None is in the heteronereid state. Some are accompanied 
by membranous tubes similar to those of N. dumerilii. 

Ehlers (4, p. 104), and Benham (1, p. 238), have indicated the wide range of this 
species in the southern oceans, and reference should be made to these papers for the 
synonymy. 

Nereis {Platynereis) dumerilii, Aud. et Edw., subsp. n. comata. (PL III. figs. 11-13.) 
Nereis {Platynereis) dumerilii, M 'In tosh (11), p. 302, and figs. 

Station 536. 27° 23' N., 33° 06' W. One example, June 28, 1904. 

Station 538. 32° 11' N., 34° 10' W. Two examples, June 30, 1904. 

These were taken in mid-Atlantic, with the tow-net, among the floating gulf-weed 
of the Sargasso Sea. Those from Station 538 were in typical membranous tubes among 
the branches of the weed. They are small specimens, about 20 to 35 mm. long. One 
at least is full of ova. 

The proboscis was in all cases inverted ; the two larger specimens were therefore 
dissected ; the proboscis of one of these, detached, flattened out and mounted in balsam, 
shows very clearly an armature of the type usual in N. dumerilii, that is, pectiniform 
series of paragnaths in the areas III., IV., VI., and VII.-VIII. The last are practically 
uniserial, and extend nearly half-way round the basal ring ; but some short combs, each 
of a few teeth, occur in front, representing the more typical biserial arrangement. 



COLLECTED BY THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 47 

The bristles are of the forms characteristic of N. dumerilii. Two or three homo- 
gomph falcigers, with free appendages, occur on the inferior margin of the notopodial 
bundle in the more posterior parapods. 

The parapods (figs. 12-13) are of normal form, except that the notopodial cirri are 
of extraordinary length, and the neuro-setigerous lobe in the round-lobed parapods 
(5th-llth pairs) is more prominent than usual. 

The noto-cirri are longest about the 15th-20th pairs, behind which they grow 
somewhat shorter. The neuro-cirri are also unusually long, distinctly overpassing the 
neuro-ligules. 

The tentacular cirri are of great length, the longest reaching approximately to the 
20th setigerous segment. 

On the head there are peculiar markings (fig. 11); a broad ring of dark brown 
granular pigment exists on the basal joint of each of the palps, and a band of similar 
nature on the forehead surrounds the bases of the tentacles. Otherwise the specimens 
— preserved in alcohol — have lost their colour. 

The extraordinary elongation of the parapodial and tentacular cirri in these 
examples seems to justify their being assigned a distinct sub-specific name. So far 
as I am aware, the only other species of nereid which shows a similar development 
of cirri is N. mirabilis, a member of quite a different group. 

Judging from the position in mid-Atlantic where these worms were collected, one 
may suppose they had drifted across from the neighbourhood of the Gulf of Mexico 
with the south-west wind drift ; but whether this variety of N. dumerilii characterised 
by such peculiarly elongate cirri has been evolved solely under the conditions of the 
drifting weed of the Sargasso Sea, or whether it is also to be found in the littoral 
areas of the seas of Central America, is a matter for future investigation. 



EXPLANATION OF PLATE III. 



Figs. 1 and 2. Nereis pelagica, Lin. South Africa. 

Fig. 1. Head and everted proboscis, dorsal view. x 18. 
,, 2. Proboscis everted, ventral view. x 18. 

Figs. 3 to 10. N. falklandica, n. sp. Falkland Islands. 

Fig. 3. Head, from above. x 15. 

,, 4. Maxillary ring of proboscis, dissected from the ventral side. x 15. 

,, 5. Basal ring, dissected from ventral side, x 15. 

,, 6. Jaw dissected out, ventral aspect. x 15. 

,, 7. 10th parapodium. x 21. 

8. 51st „ x2l. 

9. 70th „ x21. 

„ 10. Falcigerous seta, from inferior neuropodial bundle of 10th parapodium. Zeiss 4 D. 



48 MR L. N. G. RAMSAY ON POLYCHtETA OF THE FAMILY NEREIDS. 

Figs. 11 to 13. N. dumerilii, A. et E., subsp. n. comata. Sargasso Sea. 

Fig. 11. Anterior extremity from above. x 17. 
,, 12. 8th parapodium. x 32. 
„ 13. 14th „ x32. 



REFERENCES. 



(1) Benham, W. B., Report on the Polychseta of the Sub-antarctic Islands of Neiv Zealand, Wellington, 
1909. 

(2) Ehlers, E., " Florida- Anneliden," Mem. Mas. Comp. Zool. Harvard, vol. xv., 1887. 
(3) Polychdten der Hamburger Magalhaensischen Sammelreise, Hamburg, 1897. 

(4) "Die Polychiiten des magellanischen mid chilenischen Strandes," Festschr. K. Qes. Wiss. 

Gbttingen, 1901. 

(5) " Neuscelandische Anneliden," Abhandl. K. Ges. Wiss. Gbttingen, Math.-phys. Kl., N.F., iii., 

1904. 

(6) " Bodensiissigen Anneliden," Wiss. Ergebn. d. Deutschen Tiefsee-Exped., XVI. i., Jena, 1908. 

(7) Gravier, Ch., "Annelides Polychetes de la mer Rouge," Nouv. Arch. Mus. d'hist. nat., Paris, 
4° serie, ii. et iii., 1900-01. 

(8) "Annelides Polychetes recueillies a Payta (Peru)," Arch. Zool. Paris, Ser. 4, x., 1909. 

(9) Kinberg, J. G. H., "Annulata Nova," Of vers, af K. Sv. Vetensk. Akad. Fbrh., 1865. 

(10) M'Intosh, W. C, " Polychseta," Sci. Results, H.M.S. Challenger: Zool, xii., London, 1885. 

(11) The British Annelids, II., i. and ii., The Ray Society, London, 1910. 

(12) Marenzelleu, E. von, "Polychaten der Grundes-Exp. S.M.S. ' Pola,'" Denkschr. K. Akad. Wien, 
lxxiv., 1904. 

(13) Treauwell, A. L., " Polyclisetous Annelids of Porto Rico," Bull. U.S. Fish. Comm., xx. (ii.), 1900. 

(14) Malaquin, A., et Dehorne, A., "Annelides Polychetes de la Baie dAmboine," Revue Suisse Zool., 
xv., 1907. 

(15) Fauvel, P., "Annelides Polychetes du Golfe Persique," Arch. Zool. Exper. (5°), vi., 1911. 



Trans. Roy. Soc. Edin' 



Ramsay: "Scotia" Nereid.*. 



Vol. L.— Plate III 




1 l/^S^ | 



\ 
v 









wv 






10 





LN.G.R.del 



M I triune * Frckine. F.itli . KHi 



( 49 ) 



IV. — On the Genus Porponia and Related Genera, Scottish National Antarctic 
Expedition. By Professor Oskar Carlgren, Universitetets Zoologiska Institution, 
Lund. Communicated by Dr W. S. Bruce. 

(MS. received August 12, 1913. Read January 19, 1914. Issued separately March 30, 1914.) 

[Plate IV.] 

In an Appendix to the Actiniae of the Challenger Expedition, P. Hertwig. 1882, 
described a peculiar genus, Porponia, with two species, P. elongata and P. robusta, 
which he characterises in the following manner: " Actiniarien (Hexactinien?) mit 2 
Schlundrinnen ohne Ringmuskel, mit diinnwandigen Tentakeln, deren Basen auf der 
ausseren Seite durch spangenformige Verlangerungen des Mauerblatts gestiitzt werden." 
Partly owing to the badly preserved material, however, he did not venture to indicate 
definitely its systematic position, though he considered it conceivable that it repre- 
sented a transitional form between the Zoanthidse and the true Actiniae, or Hexactiniae. 
Hertwig expresses the following opinion regarding the systematic position of the 
genus : <; Die doppelreihige Stellung der Tentakel, die Abwesenheit vollstandiger 
Geschlechtsepten (Macrosepten) und unvollstandiger, sterilen Septen (Microsepten) 
sind Merkmale, welche an die Zoantheen erinnern, die Zahlen der Tentakel und der 
Septen passen ebenfalls am meisten fiir diese Gruppe, da sie weder von dem Numerus 6 
wie bei den Hexactinien noch von dem Numerus 4 wie bei den Paractinien bestimmt sind. 
Auf der anderen Seite nahert sich die P. elongata durch den Besitz von zwei Schlund- 
rinnen wieder mehr den Hexactinien, unter denen sie am meisten mit den Antheo- 
morphiden ubereinstimmt. Ich halte es daher fiir sehr wahrscheinlich, dass P. elongata 
eine Mittelform ist, welche den Ubergang von den Hexactinien zu den Zoantheen 
bildet." Porponia possibly, he thinks, belongs to the Antheomorphidse, a family 
supposed to be separated from the family Antheadse chiefly by the absence of a 
sphincter and by the weak development of the musculature. 

Since R. Hertwig described this genus, it has not been made the subject of any 
close examination, nor for this reason has its systematic position been discussed in 
detail. Yet it is only right to mention that M'Murrich was inclined to refer the 
genus Halcurias {Endoccelactis) to the neighbourhood of Porponia. " In fact," he 
writes (p. 226, 1898), " I was inclined at first to associate it (Halcurias) with Porponia, 
and was only deterred from doing so by the simplicity of the arrangement of the 
mesenteries." (That Halcurias possesses a peculiar arrangement of the mesenteries, 
which agrees with what I have described, 1897, for the genus Endoccelactis, was not 
known to M'Murrich at that time.) 

A closer examination of the material which I received for investigation from the 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 4). 7 



50 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

Scotia Expedition has proved, however, that the arrangement of the mesenteries, as 
also indeed a number of other characters, indicates a close relationship between 
Halcurias (Endoccelactis) and Porponia, though each genus has its own distinctive 
characteristics. They must of necessity be referred to the same family, though this 
cannot be the Antheomorphidse set up by Hertwig, with its genera Antheomorphe and 
llyanthopsis. As the following description will show, I have no hesitation in retaining 
the family set up by me, Endoccelactidse, for Halcurias and Porponia. 

After describing in detail the Porponia species of the Scotia Expedition, I shall 
discuss the mutual relationship of Halcurias and Porponia, and the position of the 
family Endoccelactidge in the classification. At the same time I shall take the 
opportunity to discuss the genera Antheomorphe and llyanthopsis, as also the family 
Antheomorphidse. 

I. The Structure of Porponia Antarctica, n. sr 

Place of discovery.— Off Coats Land, 71° 22' S., 16° 34' W., 1410 fathoms, 16th 
March 1904. 17 specimens. 

Dimensions of the largest specimens. — Length 6-8 cm., breadth of the foot 3-4 
cm., breadth of the disc 8-9 cm., length of the inner tentacles 3-7 cm. 

External appearance. — The fresh colour is creamy white, tinged, especially on the 
tentacles, with pale lavender. The base is expanded and often attached to a round 
stone, which it more or less encloses ; it is arranged in coarse, irregular folds, and 
secretes a fairly extensive cuticle. 

The body-wall is more or less beaker-shaped, arising from the less or greater 
contraction of the individuals. The distal part of the animal is thus wider than the 
proximal, and that often to a fairly considerable extent. The thick body-wall is 
provided with irregular longitudinal and transverse furrows, by means of which it is 
divided into irregular areas, as a rule very prominent, since the thin ectoderm has 
fallen off from almost all the specimens. The distal, uneven edge of the body-wall is 
not marked off by any definite groove or line, but passes irregularly into the bases of 
the tentacles. In large specimens the tentacles are typically 68 * in number, yet this 
may be exceeded, as the arrangement of the tentacles and even the grouping of the 
mesenteries may be somewhat irregular in one quadrant (or several ?), resulting in a 
somewhat greater development of tentacles here than in the other quadrants. The 
tentacles have the appearance characteristic for Porponia ; on the outer side they are 
greatly thickened and like cartilage ; towards the oral disc, on the inner side, they are 
thin and resemble ordinary tentacles. Owing to the mesogloea being greatly thickened 
on the outer sides of the tentacles, it looks as if the tentacles here were provided with 
bridge-like outshoots from the body-wall. Further, the tentacles are conical, curved 

* The normal number of tentacles seems to be developed in the species at a comparatively early stage, as specimens 
of only half the size of the largest in the collection have already the typical number of tentacles. A small specimen, 
on the other hand, had considerably fewer tentacles ; but as it was much damaged, I have not tried to ascertain exactly 
the number of tentacles or their arrangement. 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 51 

like a sabre towards the oral disc, thick in the basal part but gradually narrowing 
towards the tip, and of moderate length. Their arrangement is irregular, and recalls 




Fig. 1a. 




X 



<% x 



Fig. 1b. 
Fig. 1. — Porponia antarctica ; A from the side, B from the disc. 

the condition in Halcurias (Endoccelactis). In the latter genus the true arrangement 
is difficult to find, and this is even more the case in Porponia owing to the greatly 
swollen bases of the tentacles. So far as I could find, the arrangement of the tentacles 



52 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

on each side of the sagittal plane is as follows : 1 (dt.), 2, 1, 4, 3, 5, 3, 4, 1, 2, 1, 4, 3, 
5, 3, 4, I, 2, 1, 4, 3, 5, 3, 4, 1, 2, 1, 4, 3, 5, 3, 4, 1, 2 = 34 (dt. =directive tentacle). 
Altogether, therefore, there should be 18 tentacles of the first order, 10 tentacles of the 
second, 16 of the third, 16 of the fourth, and 8 of the fifth. But it has to be remarked 
that some groups of tentacles of the first and second orders, namely, the 5 (1, 2, 1, 2, 
l) in the sagittal plane on both sides of the angles of the mouth, and the 3 (1, 2, l) on 
both sides in the transverse plane, arise somewhat further in than the other tentacles 
of the first and second order. The tentacles of the first order should therefore, perhaps, 
best be divided into 2 circlets (10 + 8), and similarly those of the second order into 
2(6 + 4). The underlined tentacles of the fourth order stand somewhat further out 
than the others of the fourth order. The arrangement of the tentacles is thus 18 
(10 + 8)+ 10 (6 + 4) + 16 + 16 (8 + 8) + 8 = 68 (text-fig. 2). As in Halcurias, the 
peculiar arrangement of the tentacles is connected with a characteristic dislocation of 
the mesenteries. 

The oral disc is wide, expanded, marked by distinct radial furrows corresponding to 
the insertions of the mesenteries. The oesophagus is wide, oval, long, and provided 
with longitudinal furrows to a number of about 18. There are 2 oesophageal grooves, 
lying typically in the angles of the mouth, and broader in the aboral than in the oral 
part. No hyposulcus is developed. 

Anatomical structure. — The ectoderm of the base is high, and consists chiefly of 
supporting cells, which secrete a fairly thick cuticle. The mesoglcea is extensive, as 
also the entoderm, in which the nervous system seems to be well developed. 

The ectoderm of the body- wall by comparison with the thick mesogloea is thin and 
provided with numerous spirocysts of varying size up to 60 m long and 11m broad. 
Further, there are generally thick-walled capsules (length 26-34 n, breadth 3 m). In 
cross-section there seem to be thickenings at the base which resemble transversely cut 
muscles, but are probably in reality thickened basal parts of the ectodermal cells 
(see below under Ilyanthopsis elegans). The mesoglcea is thick, and provided with 
numerous small cells with outshoots. The entodermal musculature is weak, and no 
sphincter is present. The entoderm is thin like the ectoderm. The entoderm of the 
body-wall and of the oesophagus is pigmented. 

The ectoderm of the tentacles is somewhat high, and contains fairly numerous 
spirocysts and thick-walled nematocysts (length 36-50 m, breadth 3-(5) m). The 
longitudinal musculature on the outer side of the tentacles is weak, but gradually 
becomes considerably stronger towards the inner side, so that the muscular wall is here 
about half the height of the supporting cells. The mesogloea agrees in structure with 
that of the body- wall. On the inner side, in the lower part of the tentacles, it is, as a 
rule, just as thick as, or thicker than the ectoderm ; on the outer side, however, it is much 
thickened (fig. 6, PL IV.). The thickness decreases towards the tips of the tentacles 
(fig. 5, PI. IV.), so that at the ends the mesogloea is almost equally developed round the 
tentacles. The entoderm is thin, and the ring-musculature weak. The radial muscula- 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 53 

ture of the disc is in the ridges almost as well developed as the longitudinal musculature 
on the inner side of the tentacles, but in the furrows it is considerably feebler. 

The ectoderm of the oesophagus shows the usual structure, but in addition numerous 
thick- walled nematocysts (length 43-48 m, breadth 5 m), and in fair numbers spirocysts 
of the same appearance as in the body-wall. The mesoglcea, which is very thick, agrees 
in structure with that of the body-wall. 

The arrangement of the mesenteries is shown by the accompanying schematic 
fig. 2 ; 6 pairs of mesenteries of the first order, with 2 pairs of symmetrical, direc- 
tive mesenteries are present. While all the primary exocoels are reduced, so that no 
mesenteries could be formed in them, each of the 4 lateral endocoels contains a pair 
of mesenteries of the second order, in which, however, the longitudinal muscles face 
outwards, as in the directive mesenteries. The mesenteries of the first and second 
order, all of which are perfect, are thus 20 altogether; of these 16 are grouped 
in 8 pairs with longitudinal muscles as the directive mesenteries, owing to the develop- 
ment of the mesenteries of the third order in the secondary endocoels, and the last 4 
mesenteries are unpaired at the sides of the directive mesenteries. The arrangement of 
the 20 oldest mesenteries thus agrees completely with the corresponding condition in 
Halcurias. In the endocoels of the second order we find mesenteries of the third, 
fourth, and fifth order. The mesenteries of the third order (8 pairs) are perfect, and 
have longitudinal muscles typical of the Actiniaria (muscles facing inwards). The 
mesenteries of the fourth order are unequally developed. On one side of the 
mesenteries of the third order, between these and the mesenteries of the second 
order, there is an imperfect mesentery of the fourth order, but on the other side a 
pair, consisting of one imperfect mesentery, lying nearest the mesenteries of the 
third order, and a perfect mesentery. Between the latter and the mesentery of the 
first order there is an imperfect mesentery of the fifth order. The arrangement of 
the mesenteries is thus : 6 — 4 — 8 — 8 + 8 unpaired — 8 unpaired ; thus in all 26 pairs, 
+ 8 unpaired of the fourth and 8 unpaired of the fifth order, or 68 mesenteries. 
While this is typical, the development probably proceeds somewhat further in some 
ways in very old specimens. 

The mesenteries are somewhat thick, owing to the thickness of the mesoglcea. The 
longitudinal musculature is fairly well developed, though not so much as in Halcurias. 
In the distal part the folds are thick, but grade towards the proximal end to a weak, only 
a little condensed, pennon-like region (figs. 7, 8, PL IV.), which approaches the body- wall 
and fuses with the well-developed parietal parts of the longitudinal musculature. The 
parietobasilar musculature is narrow but fairly well developed (figs. 7, 8, PL IV.), and 
goes far out towards the distal end. There are no separate basilar muscles, though in 
the foot end the muscles pass in a transverse direction, also on the longitudinal muscles' 
side of the mesenteries ; but these muscles which run transversely are continuations of 
the longitudinal musculature, which at the base of the column bend in a transverse 
direction (cf. p. 60, Ilyanthopsis elegans). 



54 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

The filaments are well developed and extremely broad, owing to the strong develop- 
ment of the mesogloea (fig. 9, PI. IV.). The vacuolar streak is little differentiated. Both 
in the intermediate part of the ciliated tract region and in the nematocyst glandular 
streak there are sparse spirocysts and fairly numerous thick-walled nematocysts, 
especially in the latter (length 36-41, sometimes even 46 m, and about 3 m broad). Of 
fairly common occurrence further in the glandular streak are nematocysts with distinct 
basal part to the spiral thread (length 37-41 m, breadth 7 m). Even the entoderm of 
the filament is pigmented, especially on the region of the border-streak. 

The sexual organs occur on all well-developed mesenteries, even on the directive 
mesenteries. The animals are dioecious. 

For the Porponia obtained by the Scotia Expedition I have set up a new species, 
P. antarctica. Of the species of Porponia already known it comes nearest to 
P. robusta, R. Hertwig, both in the form of the body and the appearance of the 
tentacles. To set up good characters between these two species is, however, distinctly 
difficult, as Hertwig's description of P. robusta is so incomplete, and both, this species 
as well as the two specimens of P. elongata, do not seem to have the number of 
mesenteries and tentacles typical of P. antarctica. It is probable that the mesenteries 
of the fifth order have not been laid down in Hertwig's form, to judge from the 
number of tentacles, which Hertwig gives to be 54 in P. elongata. According to 
some notes made by me in 1897 on revising the Challenger Actinias, the distribution and 
size of the spirocysts and nematocysts in P. robusta and P. elongata were as follows : — 

The spirocysts in the body-wall were in P. robusta very numerous and about 
40-44 m long, in the tentacles of P. elongata about 56-72 m ; there were also spirocysts 
in the oesophagus of these two species. The nematocysts in the oesophagus were 48 m 
long in both P. elongata and P. robusta, in the tentacles of P. elongata 48 m long. 
It is, however, noticeable that there were only fragments of the ectoderm in the 
Challenger species. 

II. On the Systematic Position op the Genus Porponia. 

According to the anatomical account of the genus Porponia given above, there 
should be no doubt remaining that Porponia and Halcurias (JSndocoelactis) are very 
nearly related to each other. Common to both is the structure of the body-wall, and 
also of the oesophagus, both, among other things, being provided with numerous spiro- 
cysts. Even the anatomical structure of the filaments and distribution of the sexual 
organs show agreement. Most striking, however, is the characteristic and similar 
arrangement of the tentacles and mesenteries, which differs from that in all other 
known Actiniaria, as can be seen more clearly from the following scheme for the two 
genera. In both Halcurias and Porponia the same displacement of the original 
mesenteries and tentacles has clearly taken place. After the formation of the first 6 pairs 
of mesenteries, 1,1, etc., in the ordinary way, 2 mesenteries (2, 2), with the longitudinal 
muscles faced outwards, have arisen in the lateral endocoels, and these mesenteries 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 55 

form normal pairs with neighbouring mesenteries of the first order. At this stage, 
therefore, the genera have 10 pairs of mesenteries, 2 of which, lying medially 
opposite to each other, are directive mesenteries, whilst the other 8 are formed in 
pairs with typically arranged longitudinal muscles. In the 8 new endoccels arisen 
through the growth of the mesenteries of the second order, there is a further develop- 
ment of 8 normal pairs of the third order, so that at this stage we have 10 (6 + 4) + 8 
= 18 pairs of mesenteries. Thereafter the development proceeds in a more normal 
manner, the mesenteries of the fourth and fifth order in Porponia not developing in 
the endocoels of the third order, but on both sides of a pair of the third order ; in an 
exocoel in relation to the mesenteries of the third order, but in an original endoccel of 
the second order. I have endeavoured previously (1897) to show that the develop- 
ment proceeds in this way in Endoccelactis, as with fairly great certainty I considered 
it possible, partly from the unequal development of the mesenteries at the basis of the 
column, to distinguish the first 6 pairs of mesenteries (1,1, etc.) from the 4 pairs of the 
second order, and thence from the obviously great development of these 10 pairs in 
comparison with the others to distinguish them clearly from the subsequent orders of 
mesenteries. In Porponia I had admittedly not been able to study the development 
of the mesenteries so clearly as Endoccelactis, as the difference between the mesenteries 
of the said orders were not so distinct as in this latter form, owing to the fact that 
in Porponia several more complete mesenteries occur than in Endoccelactis. The 
arrangement of tentacles in Porponia, as also the whole arrangement of mesenteries, 
indicates, however, that in this genus the mesenteries arise in the same manner, and that 
also after reaching a stage with 6 normally placed pairs of mesenteries, a development 
of mesenteries in the endocoels has taken place. The groups of tentacles of the first and 
second orders, which arise near the directive mesenteries 1, 2, 1 (dt.), 2, 1, and those 
which lie in the transverse plane 1,2, 1, are inside the tentacles of the corresponding 
order which stand at the other 4 pairs of stronger mesenteries, a condition that is to 
some extent indicated on the schematic figure, but which in reality is considerably 
greater than the figure shows. This indicates that we have to arrange the first 6 pairs 
of mesenteries by this plane. With regard to the arrangement of the tentacles other- 
wise, this is in the main the same in the two genera, great displacements occurring in 
the cycles with the development of certain mesenteries in the endoccels, wherewith, so 
to speak, a doubling of the tentacles in the lateral endocoels of the first and second 
orders arises. Above all, the arrangement of the 28 innermost tentacles is the same 
in both genera, as the figure shows. It is characteristic of both genera, therefore, that 
all the mesenteries of the second and third orders develop in the endoccels, and that 
in consequence great displacements in the position of the tentacles take place. 

Nevertheless, there are a number of differences in the structure of the two genera. 
In a number of less important characters, such as the form of the body — in Porponia 
beaker-like, in Halcurias more cylindric — in the presence of only one oesophageal 
groove in Halcurias, whilst Porponia has two, there are certainly differences between 



56 



PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 




Fig. 2. 




Fig. 3. 

Figs. 2 and 3. — Schematic sections through the oesophagus region of Porponia antarctica (fig. 2) and 
Halcurias rarlgrcni (fig. 3). The position of the tentacles (the dark rings) is drawn in. The rect- 
angular, dark figures on the mesenteries indicate the longitudinal muscles. The shaded liDes denote 
the directive plane. In Porponia the ridges of the oesophagus are not drawn in. 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 57 

the genera, but also others more important exist. In the two genera the tentacles are 
formed in a different manner, having the usual appearance in Halcurias, whilst they 
are greatly thickened in Porponia on the outer side, so that it looks as if the thick 
mesoglofta of the body-wall extended in the form of bridge-like outshoots into the 
tentacles. An important distinction between Porponia and Halcurias is also seen in 
the development of the mesenteries. Whilst Halcurias has only 10 pairs of stronger 
mesenteries — the mesenteries of the third order being thus but feebly developed, and 
that only in the most distal end — there is a much larger number of stronger mesenteries 
in Porponia, which are connected with the oesophagus in its whole length or the 
greater part of this. Very characteristic also is the fact that the development of the 
mesenteries, which appear after the mesenteries of the third order, is different in the 
two genera. In Halcurias the mesenteries of the fourth order arise normally, so that 
the two mesenteries in the same pair appear simultaneously ; the development then 
ceases, though it is possible that in some cases mesenteries of a fifth order may be laid 
down. In Porponia, on the other hand, the development has taken a different line, 
and comes to resemble that of the later mesenteries in Actinostola and Stomphia. The 
mesenteries of the fourth order, namely, do not appear at the same time ; on the one 
hand, the one mesentery is developed much earlier than its pair ; on the other, the 
development of the mesenteries of the fourth order is delayed in certain regularly 
arranged exocoels, so that here only one unpaired mesentery instead of a pair arises. 
This displacement of the mesenterial appearance has had the result that an unpaired 
mesentery of the fifth order arises on the side where the strongest and perfect mesentery 
of the fourth order lies. In fact, the development of the mesenteries of the fourth and 
following orders in Porponia comes under the same law as I have indicated for the 
development of later mesenteries in Actinostolidse, and which means that the develop- 
ment of a stronger mesentery in a previous order leads to an earlier development 
of the mesenteries of the subsequent orders in the area which lies nearest this 
stronger mesentery. Finally, it has to be noted that irregularities not rarely arise in a 
quadrant of Porponia which shows a somewhat larger number of mesenteries than the 
normal. If we indicate the mesenteries with numbers, those of the first order with 1, 
of the second with 2, and so on, the irregularity in the arrangement of the mesenteries 
on each side of the directive plane can be seen from the following scheme for the two 
genera (dm. = directive mesenteries) : — 
Halcurias {Endocadactis). 

1 (dm.) 1, 4, 4, 3, 3, 4, 4, 2, 2, 4, 4, 3, 3, 4, 4, 1, 1, 4, 4, 3, 3, 4, 4, 2, 2, 4, 4, 3, 

3, 4, 4, 1, 1 (dm.) = 34. 
Porponia. 

1 (dm.) 1, 5, 4, 4, 3, 3, 4, 2, 2, 4, 3, 3, 4, 4, 5, 1, 1, 5, 4, 4, 3, 3, 4, 2, 2, 4, 3, 3, 

4, 4, 5, 1, 1 (dm.) = 34. 

The arrangement of the tentacles is also somewhat different in the two genera (see 
figures). This difference is due entirely to the different arrangement of the mesenteries 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 4). 8 



58 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

of the later developmental stages, yet it has to be noted that the position of the 
tentacles, especially in Porponia, is difficult to determine, and it is just possible that 
the agreement is somewhat greater than is represented in the schematic figures. 

While Halcurias and Porponia are thus closely related to each other, there is still 
the question whether they have any near relationship to the genus set up by R. Hertwig, 
Antheomorphe, as M'Murrich and Hertwig maintain, the one for Halcurias, the other 
for Porponia. In my opinion, such a relationship does not exist, for Antheomorphe, 
according to Hertwig's poor description of this genus, appears to have normally arranged 
mesenteries and tentacles. On the other hand, it is not impossible that Halcurias and 
Porponia are related to the form which Wassilieff (1908) has described as Ilyanthopsis 
elegans. The abnormal development of the mesenteries in Ilyanthopsis elegans, ac- 
cording to the obviously imperfect data given by Wassilieff, indicates, namely, the 
possibility of a mesenterial arrangement such as in Halcurias and Porponia, and even 
the habitus of the animal resembles that of Halcurias. To settle this, I have obtained 
a specimen of Ilyanthopsis elegans for investigation from the Conservator of the Bavarian 
State's collection in Munich. As Wassilieff's description of the animal leaves much to 
be desired,* and as it is by no means so schematically constructed as this author believed, 
I give here a description of this species, a description, however, that cannot be considered 
complete, as the material, which could not be dissected, was insufficient for the purpose. 

The body in Ilyanthopsis elegans is elongated, cylindrical, the base distinctly 
flattened, but the boundary between body-wall and base not sharply marked. The 
body-wall is provided with irregular transverse and longitudinal furrows, so that the 
parts of the wall between the furrows have the appearance of raised areas, which, how- 
ever, do not differ in their structure from the remaining part of the body-wall. The 
tentacles are of moderate length, 2-2*5 cm. (length of body over 6 cm.), and distinctly 
furrowed longitudinally, not thickened at the base and gradually narrowing towards 
the distal end. The arrangement of the tentacles was difficult to determine and by no 
means so simple as Wassilieff imagined. There is no arrangement into two circlets 
only ; on the contrary, the arrangement shows distinctly a certain resemblance to that 
in Halcurias and Porponia, as some displacement of the tentacles has taken place, 
here also assuredly connected with an irregular arrangement of the mesenteries. In the 
first place, we have in each angle of the mouth (in the sagittal plane) quite the same 
arrangement of the tentacles as in Porponia and Halcurias. They further agree in 
this, that at certain points groups of tentacles occur, where two tentacles of the first 
order stand on each side of one of the second order. Lastly, the other tentacles also 
show a resemblance in arrangement to the corresponding tentacles in these genera. 

* The Actiniaria described by Wassilieff, especially some of the species, require revision. Thus, the Actinia 
described by Wassilieff under the name of Halcampella minuta is quite certainly no Halcampella, but rather a 
Haloclava. The occurrence of warts and the appearance and structure of the tentacles, which are bulb-like and swollen 
at the tips, speak in favour of this. The arrangement of the mesenteries and the siphonoglyphe agree less definitely, 
but this may be due to the specimen being a young individual, as is indicated by the small number of tentacles (15). 
If it cannot be referred to Haloclava or Eloactis, it may represent the type of a new genus. 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 59 

The arrangement, however, is difficult to determine in detail in Ilyanthopsis, as the 
grouping of the mesenteries is not known, and only one specimen has been at my 
disposal, which, moreover, showed an irregular grouping of the tentacles in certain parts. 
Starting from the one directive tentacle (that where the directive line passes through 
an oesophageal groove), the arrangement seems to be the following (l, 2, 3, etc., tentacles 
of the first, second, third, etc., order ; dt. =the directive tentacle) : — 

1 (dt.)-2-l-5, 4-5-4-5-3-4-3-5-4-5-4-5-1-2-1-5-4-5-4-5-4-5-1-2-1-5-4-5- 
4-5-4-5-1-2-1-5-4-5-4-5-4-5-1-2-1- (dt.)-2-l-5-4-5-4-5-4-5-l-2-l-5-4-5-4- 
5-4-5-4-5-1-2-1-5-4-5-4-5-4-5-1-2-1-5-4-5-4-5-4-5-1-2. 

It is to be noted here, that among the last mesenteries of the first order, nearest to 
the last mesentery of the first order, there may be a few tentacles more. A small part 
of the edge of the body, namely, had been torn away, and it is possible that some 
tentacles, at most 4 probably, have gone with it. In this specimen, therefore, the 
number of tentacles is somewhat higher, yet not so high as 100. The group 3, 4, 3, is 
also somewhat uncertain, in so far as it cannot be determined whether it is a 1 , 2, 1 
group or not ; in any case it seems to stand somewhat inside the other groups of the 
first and second order. But to clear up the arrangement of the tentacles definitely 
requires an investigation of more material with the developmental forms. 

The oral disc is somewhat broader than the base, with deep, radial furrows corre- 
sponding to the insertions of the mesenteries. The mouth was partly projecting, wide. 
The oesophagus is wide, long, almost half the length of the body, with a siphonoglyphe. 
Wassilieff states that two grooves are present, but I have not been able to find more 
than one ; the other supposed to be present by Wassilieff does not differ — at least in 
the specimen examined by me, which has also been investigated by Wassilieff — from 
the deep longitudinal furrows with which the oesophagus is also provided. Including 
the siphonoglyphe, the number of these furrows amounts to about 16. No well-marked 
prolongation of the groove downwards below the oesophagus is present. 

The anatomical structure resembles that of the Endocoelactidse. The ectoderm of 
the body-wall is not extensive, especially by comparison with the thick mesogloea. It 
contains very numerous spirocysts of varying size, and fairly numerous thick-walled 
nematocysts (length 34-43 m, breadth 5 n). In transverse sections at the base of the 
ectoderm we find structures packed close together, which greatly resemble cross-sections 
of ectodermal, longitudinal muscles. In the beginning also I was inclined to take 
them as such, but a closer examination showed that this was hardly the case. In 
longitudinal sections (fig. 2, PI. IV.) I found similar structures, which should not have 
been there if it was a question of longitudinal muscles. The muscle-like parts are, 
indeed, so far as I could determine, no other than the greatly thickened bases of 
ectoderm cells, which can also be clearly seen when the section cuts through a wall of 
the ectoderm and passes through the ectoderm cells a little way from the base. In 
these it can be seen that the ectoderm cells are fixed in the mesogloea by greatly 
thickened bases. I lay stress on this, as M'Murrich was of the opinion that in 



60 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPOMA 

Halcurias pilatus he had found longitudinal muscles in the ectoderm of the body-wall. 
Though I have not examined this species, I consider it possible that the same condition 
prevails there as in Ilyanthopsis — in other words, that the longitudinal muscles are 
no other than thickened, basal parts of the ectoderm cells. The mesogloea of the 
body-wall is very thick, and provided with extremely numerous cells with bipolar, 
irregularly prolonged ends. In the inmost part of the mesogloea there are closely 
packed fibrillar folds arranged as fig. 1, PL IV. shows. The sphincter I have not 
closely examined, but I think it probable, from an external investigation, that this is 
completely wanting, as Wassilieff states. The entoderm is somewhat broader than 
the ectoderm, and thicker in the middle between the insertions of the mesenteries than 
at the sides. Wassilieff has described the structure of the tentacles and emphasised 
the longitudinal muscles of the ectoderm. The ectoderm contains very numerous 
spirocysts of varying size, and also numerous thick-walled nematocysts (length about 
36 m). The longitudinal musculature of the ectoderm is comparatively weak. How 
far fine outshoots run out from the mesogloea into the ectoderm, as is stated by 
Wassilieff, I am unable to determine ; it seems to me that these outshoots are 
nothing else but the thread-like basal parts of the ectoderm cells. The ectodermal 
nematocysts and spirocysts of the oral disc agree with those in the tentacles, but are 
not so abundant. The ganglion and nerve layer is also well developed here, as in the 
tentacles. The ectodermal radial musculature is strong, especially in the ridges, and 
the muscle folds just as high as, or higher than, the epithelial parts of the ectoderm. 
The inner parts of the muscle folds show a tendency to be mesogloeal in the ridges 
(fig. 3, PL IV.). The structure of the oesophagus agrees with that of the body- wall. 
The mesenteries are all complete, corresponding in number to the tentacles. 

As Wassilieff states, the musculature is very weak, and only somewhat more 
strongly developed where the mesenteries join the body-wall (fig. 1, PL IV.). Here we 
can distinctly distinguish the muscles, both the longitudinal and the parieto-basilar, 
which are weak. On the very thin mesenteries there is no trace of protuberance of 
the longitudinal muscles. In longitudinal sections through the mesenteries and trans- 
verse sections through the base it looks as if the basilar muscle occurred as a weak 
fold of the musculature of the mesenteries. But specially differentiated basilar muscles 
do not seem to be developed, for the transverse layer of muscles near the base on 
the side of the mesenteries where the longitudinal muscles are, is formed by 
the latter muscles, bending almost at a right angle a little way from the base, 
thus forming what may be called pseudo-basilar muscles (fig. 7, PL IV.), the same 
condition as in Porponia, as I was able to determine from preparations of the 
mesentery. With regard to the arrangement of the mesenteries, it is impossible to 
determine whether all of them are equally developed. So much can be said, however, 
that in the specimen I have examined, in addition to the directive mesenteries with 
longitudinal muscles on the outer side, the other mesenteries seem to be arranged in 
pairs with the longitudinal muscles on the inner side. The Halcurias stage, or rather 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 61 

the Porponia stage, which Ilyanthopsis probably passes through during development, 
is thus not apparent in the older individuals. The filaments seem to agree with those 
in Halcurias and Porponia, but were so badly preserved that I could not obtain any 
clear picture of them. In addition to spirocysts there are thick-walled nematocysts 
like those in the body-wall (length 36 m), as also numerous thick-walled capsules with 
distinct base to the spiral thread (length 31-34 m, greatest breadth 5 m). Sexual 
organs are present on all the mesenteries. The animal is hermaphrodite. Well- 
developed testes occurred distally inside the filament region in each mesentery, while 
a few grape-like eggs were found in the proximal part. 

The investigation of Ilyanthopsis elegans has thus led to the result I imagined it 
would, namely, that this is nearly related to Halcurias and Porponia. Though the 
material is too small to permit of a detailed statement of the grouping of the tentacles 
and mesenteries, there can be no doubt that they should be placed together. The 
arrangement of the tentacles shows the same characteristics as in these genera, and 
certain features of the mesenterial arrangement are the same apparently. Even the 
external appearance agrees well with that of Halcurias : spirocysts occur in the body- 
wall and oesophagus, as in the latter genus and Porponia. Ilyanthopsis shows most 
resemblance to Halcurias, and it might be a question whether these two genera should 
not be joined as one. For the time being, however, such a grouping would not be so 
fortunate, as Ilyanthopsis has a much greater number of mesenteries than Halcurias . 
further, in the former all the mesenteries are perfect, while in the latter about half are 
perfect. Add to this that the longitudinal musculature of the mesenteries is strongly 
developed in Halcurias, very weak in Ilyanthopsis, and it is evident that Ilyanthopsis 
has its own developmental characteristics. It seems, moreover, more probable that 
Ilyanthopsis has passed through a Porponia stage than a Halcurias stage, if the 
mesenteries are taken into consideration. If we imagine all the mesenteries in Porponia 
to be perfect, it is quite easy from them to derive the arrangement of the mesenteries 
in Ilyanthopsis. In Halcurias, on the other hand, the stronger, not-directive mesen- 
teries occur as unpaired mesenteries. How the development has proceeded we can 
only learn from the younger stages. I consider it advisable, therefore, to set up a 
new genus, Synhalcurias, for the species Ilyanthopsis elegans. The genus Ilyanthopsis 
must be abolished, as the type species of this genus, Ilyanthopsis longifilis, E. Hertwig, 
is no other than Condylactis passiflora, as stated by Pax (1910) ; I had also come to 
this view in 1897 on examination of type specimens of the Challenger Actiniae in 
London. 

We know of one more genus that might possibly be allied to Porponia, namely, 
the genus Actinernus, founded by Verrill. From R. Hertwig's description of 
Polysiphonia tuberosa ( = Actinernus tuberosus M'Murrich) and from M'Murrich's 
description of A. plebeius, however, we can hardly conclude that a close relationship 
exists between Porponia and these forms. According to my observations on a 
specimen of Polysiphonia tuberosa from the Challenger Expedition, the arrangement 



62 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

of the tentacles does not seem to show the displacement seen in Porponia ; in fact, 
as far as I can see, the tentacles are at no places grouped in such a way that two 
tentacles of the first order border on a tentacle of the second order, even though 
certain changes in the size of the tentacles have been observed, so that accord- 
ing to R Hertwig the exoccel-tentacles are not the smallest in size. Though 
Actinernus plebeius and A. tuberosus do not suggest any close relationship to 
Porponia, it is yet not impossible that the type specimen of the Actinernus genus 
may show greater similarities, a question I may leave unsettled at present, as I have 
not had the opportunity to examine this specimen.* 

For the present, therefore, we must be content with a comparison between 
Halcurias, Porponia, and Synhalcurias. The question is now, where we are to place 
these genera, and would it be of advantage to separate them from other forms 1 
M'Murrich in 1901 dealt with this question with regard to the genus Halcurias. 
" There are, apparently, three courses open for the disposal of the genus. It may be 
referred to a family already existent, the definition of the family being changed, if 
necessary, to accommodate it, or it may be taken as the type of a distinct family, as 
Carlgren has done, or, finally, it may be separated altogether from the Hexactinise 
and regarded as the type of a separate tribe. It seems to me that this last procedure 
is quite unnecessary and would probably be entirely out of harmony with the phylo- 
genetic relationship of the genus. We have learned within recent years how extensively 
nearly allied forms may differ, and how great all the modification which the hexactinian 
type may undergo. The entire facies of Halcurias is that of an hexactinian." I am 
entirely in agreement with the above citation from M'Murrich, and, like this author, 
I am of opinion that it is unnecessary to set up a separate tribe for this genus and 
Porponia, as the whole development indicates that the initial stage is a typical 
hexactinian with six pairs of mesenteries. M'Murrich comes further to the conclusion 
that Halcurias need not be placed either in a separate family, as 1 had done in 1897, 
but considers it preferable to refer the genus to the family Actinidse (Antheadse). 
"The peculiar mode of development of the secondary and tertiary mesenteries is of 
minor importance, and I see no more reason for separating Halcurias as the type of 
a new family than I do for separating an octamerous sagartian or one with a multi- 
plicity of mouths and many siphonoglyphs from the rest of the members of that family." 
He supports this view because "occasional endocoelous development of mesenteries 
have been already recorded, as in Bunodes thallia, in Actinioides dixoniana and 
papuensis" — a condition already pointed out by me in 1897. 

But is this view of M'Murrich justifiable ? So far as I can understand, this is not 
the case, as variations irregularly arisen through asexual propagation, or through 
regeneration and regulation in the symmetry of certain species — in the case of the 
phylogeny — cannot directly be compared with similar variations from the normal type 
arising during the ontogeny — a condition not hitherto taken into consideration, but 

* Compare Appendix ! 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 63 

which I must strongly emphasise. A species, for example, that normally, through 
unequal development of mesenteries, through stopping of the growth of certain parts 
and more rapid growth of others during the ontogenic development, e.g. from being 
a 6-rayed becomes an 8- or 10-rayed type, which is constant or nearly so for the 
species, cannot in phylogenetic respects be compared with another species where the 
same stages are obtained through accidental, asexual propagation or by regeneration. 
In the first case, the 8- or 10-rayed type is constant for the species, and occurs onto- 
genetically and phylogenetically ; in the latter case, on the other hand, it is a mode of 
adaptation in a less or greater part of the individual, and is dependent on the course of 
the asexual propagation, and the greater or less reduction of the old mesenteries in the 
separated or damaged fragments, a condition which has been further dealt with in my 
studies on the regeneration and regulation stages in the Actiniae, 1904, 1909. In so 
far as it has arisen ontogenetically, an 8- or 10-mesentery stage is thus of direct use for 
the phylogeny, but not in other cases. What applies to the occurrence of an 8- or 10- 
rayed type of Actiniaria also applies to the varieties that arise through the develop- 
ment of the mesenteries in the endocoels. In such cases the conditions are in full 
agreement with those found in 8- or 10-rayed forms. Porponia and Halcurias leave 
no doubt that the regular development of mesenteries in the endocoels has taken place 
ontogenetically, whereas the irregular and chance development of mesenteries in the 
endocoels in Bunodes, Actinioides, and others stands in intimate connection with the 
regeneration or possible early dislocations of tissues during development. In Porponia 
and Halcurias the development of mesenteries in the endocoels is of importance for the 
classification, whereas the abnormal occurrence of mesenteries in the endocoels in 
Bunodes, etc., is of no use for this purpose. 

The peculiarity that mesenteries occur regularly in the endocoels during the course 
of development is thus quite specific for Porponia and Halcurias, and probably also 
for Synhalcurias, and has not been observed in other Actiniaria. The question is still 
left open, if this peculiarity is of such great importance that it necessitates the setting 
up of a separate family. As mentioned above, M'Murrich connects the development 
of mesenteries in the endocoels with the occurrence of an 8-rayed type, with the develop- 
ment of several mouths and several siphonoglyphs. Just as little as we separate the 
forms showing such variations from the normal Actiniaria type ought we, in his opinion, 
to separate Halcurias from allied forms on account of the development of mesenteries 
in the endocoels. That the multiplication of mouths in a genus of Actiniaria does not 
involve a separation of the genus in question from other closely related species is 
evident from the above, as this multiplication has not arisen ontogenetically, but by 
asexual propagation. The same is certainly also the case with the multiplication of the 
siphonoglyphs. It is now left to take into consideration the abnormal development of 
the mesenteries. An 8- or 10-rayed type derived ontogenetically from a 6-rayed one is, 
as already known, by no means a seldom occurrence within the Actiniaria group, and 
may obviously arise within different families and genera that are in no genetic connec- 



64 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

tion with each other. M'Murrich is therefore quite right in saying that a genus or 
species need not be separated from other genera or species because it has been trans- 
formed into an 8- or 10-rayed type. It must be pointed out, however, that such a type 
may in certain cases be of great importance for the classification, namely, in cases where 
8 or 10 rays are observed in all species of a certain genus, as the variation in the 
symmetry can naturally be used as a good generic character. We know of no case 
where a number of the pairs of mesenteries differing from 6 has led to the setting up of 
a separate family. 

As shown above, both Porponia and Halcurias, from an assumed typical 6-paired 
mesentery stage, are transformed into one having 10 pairs of mesenteries. Where the 
transformation takes place in the ordinary manner by the belated appearance of certain 
mesenteries in certain areas and through the arising of other mesenteries in the 
exoccels,* it seems unnecessary to separate these genera, but as the 10-rayed condition 
arises in such a specific way by development of mesenteries in the endocoels, a develop- 
ment that is continued during -the following cycle, I consider it absolutely necessary to 
set up a separate family for these genera, the more so as such an ontogenetic develop- 
ment of mesenteries in the endocoels has not been observed in any other Actiniaria of a 
higher type. As far as we know, no such displacement of the tentacles has been 
observed in other forms of Actiniaria than the above mentioned. I place Porponia and 
Halcurias together in one family, therefore, to which already in 1897 I gave the 
appropriate name of Endoccelactidse. 

III. Relationship of the Family Endoccelactidse to other 
Actiniaria — Origin of the Rugosa Type. 

As already mentioned in the introduction, R. Hertwig stated the possibility that 
Porponia, owing to the arrangement of the macro- and micro-mesenteries, might form a 
transitional stage between the Hexactiniaria (Actiniaria) and Zoanthidse (Zoantharia). 
This explanation of the position of Porponia and the family Endoccelactidae cannot, 
of course, be maintained, after we have ascertained the facts on which the relation- 
ship between stronger and weaker mesenteries depends. There is nothing in the 
organisation of the family Endoccelactidee that might indicate a close relation to the 
Zoanthidae, as the development of the mesenteries in this family takes place in quite a 
different way from that in the latter characteristic group of Anthozoa. 

In my paper on Endoccelactis ( = Halcurias) I pointed out that in Minyas there is 
a strong tendency to widen the endocoels at the expense of the exocoels, causing an 
alteration in the grouping of the mesenteries, which had some resemblance to the 
alteration in the grouping of the 10 stronger mesenteries in Endoccelactis. How this 
grouping of the mesenteries has taken place in Minyas is still unknown, but it may 
possibly have arisen in connection with the development of mesenteries in the endoccels, 

* It is also to be noted that not all 8- or 10-rayed types are homologous with each other, for the 8- or 10-rayed 
condition is not always obtained in the same way ontogenetically. 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 65 

though this is not absolutely necessary. The arrangement of mesenteries in Minyas 
may be explained quite simply through an enlargement of the endoccels. In any case 
I consider the similarity in the Endoccelactis and Minyas arrangement as due to con- 
vergence, a view which I am now able to further confirm, as Minyas, i.e. the species 
described by me in 1895, and a closely related species, probably M. olivacea, later 
examined by me, are stichodactyline Actiniaria, which are nearly allied to the family 
Aurelianidse (the genera Aureliana and Actinoporus). 

With regard to the position of the family Endocoelactidse, I pointed out in 1897 
that it must be placed fairly low in the system of Actiniaria, a view that has also been 
taken up by M'Murrich. This is indicated not only by the absence of the sphincter 
and the presence of spirocysts (thin-walled nematocysts) in the ectoderm of the body- 
wall and oesophagus, but also by the absence of true differentiated basilar muscles. 
Thus, the Endoccelactids must be Actiniaria, though they are not developed in the same 
way as the elongated genera provided with physa (e.g. Edwardsidse, Halcampidse). 
From a theoretical point of view we must assume the occurrence of forms which con- 
stitute a link between the Protactininse and the Athenaria among the Actininse, i.e. 
we must take for granted the occurrence of original Actiniaria, which by the retention 
of the original body-shape with flat base (thus without development of a physa) have 
lost the longitudinal muscles in the body-wall, but, on the other hand, have not yet 
developed true basilar muscles; in the same way as I pointed out (1900, p. 57) that 
the family Discosomidse forms a link between the Protostichodactylinse and Stichodo- 
dactylinse. Among the Actininse type similar conditions would then prevail with 
regard to the family Endocoelactis, if my supposition that this family has no longi- 
tudinal muscles in the body-wall proves to be correct. Should it be the case, on the 
other hand, that M'Murrich is right in saying that such longitudinal muscles occur 
in Halcurias pilotus, the family Endocoelactidse must be referred to the Protactininse. 
In this case the thickenings of the basal parts of the ectodermal cells in the body-wall 
may be considered as rudimentary epithelial muscles, a view, however, I do not hold, 
and a question that can only be answered by means of good material of maceration. 
For practical reasons it would possibly be advisable in future to combine the family 
Endocoelactidse with the Protactininse, and the Discosomidse with the Protostichodac- 
tylinse, a grouping which I already, in 1900, p. 57 (77), pointed out as possible with 
regard to the Discosomidse. 

In my opinion, the family Endocoelactidse must thus belong to the lowest Actininse, 
or possibly to the more differentiated Protactininse. Probably an intimate relation to 
any other Actininse family does not exist. 

Before concluding the account of the relations of the family Endocoelactidse to 
other Actiniaria, we might just point out that these variations are of importance for 
the study of the other Anthozoa. As already known, the skeleton-forming Madreporaria 
show similar variations from the usual symmetry, as the Actiniaria, as 8- and 10-rayed, 
radial or more bilateral forms are found even there. As we have seen that such a 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 4). 9 



66 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

peculiar symmetry as that in Endocoelactidae may also be found in the free Actiniaria, 
it seems reasonable to conclude that among the variously attached Madreporaria, with 
their varying adaptation to the under-layer, still more complicated and varying 
arrangement of the mesenteries may be found. In my opinion, the arrangement of the 
mesenteries in Endocoelactidae opens up the possibility of a more intimate connection 
between Rugosa and Madreporaria, and more readily leads to an explanation of the 
conditions of symmetry in Rugosa, like the one proposed by me in Bronn's Klassen 
und Ordnungen, p. 1 50. Whilst the development of the mesenteries in the Endocoe- 
lactidse gives a greater possibility of interpreting Rugosa, it makes the question still 
more complicated, as in Rugosa there might be a development of mesenteries in the 
endocoels. Though we shall naturally never be able to reach finality with regard to 
the position of Rugosa as compared with the typical Madreporaria, but have to be satis- 
fied with a hypothetical explanation, so long as we do not know how the mesenterial 
musculature is arranged, I shall nevertheless give a picture of the way in which we 
might imagine the origin of the Rugosa type, if the mesenteries after the 6-pair stage 
have developed in the endocoels. I presuppose that the hypothetical, separating walls, 
sarcosepta, are taken as mesenteries, the skeletal dissepiments, sclerosepta, as septa. 

We start, therefore, from a stage with 6 pairs of mesenteries arranged typically, but 
with the lateral endocoels larger or at least as large as the exocoels. In each of the 6 
endocoels a septum has been formed (text-fig. 4a). In the next stage the development 
of mesenteries of the second order takes place in the same way as in the Endocoelactidae, 
i.e. in the lateral endocoels, 4 pairs of the second order with the longitudinal muscles 
turned outwards. Each of these mesenteries of the second order forms a new pair with 
neighbouring mesenteries of the first order. In these new endocoels 4 septa are formed 
(text-fig. 4b). 

Owing to this arrangement of the mesenteries and their occurrence only in the 
lateral endocoels, 4 zones of development have arisen instead of the 6 found in the 
exocoels of a normal Madreporaria. These zones of development lie one in each 
quadrant of the animal. This results in an asymmetrical development of the 
mesenteries, together with an irregular growth of the walls, due to the fact that the 
animal is generally attached along the one side of the goblet, or at least has been so 
once. The consequence is now that in each quadrant of the dorsal side of the animal, 
i.e. the side turned away from the siphonoglyphe, a complete suppression of the 
mesenteries of the next order takes place, while the development in the ventral part is 
continued. In the ventral endocoels 4 pairs of mesenteries arise with the same arrange- 
ment of the musculature as those of the second order. These mesenteries form new 
pairs with adjacent mesenteries of the first and second order. In the 4 new endocoels 
4 septa are formed (text-fig. 4c). The development is continued in this way with the 
next order, with suppression of the mesenteries and septa in the dorsal endocoels of the 
third order in each quadrant. At the end of the development, or at least at a late 
stage, septa develop in the exocoels (text-fig. 4d). 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 67 



As we see, the development of the Kugosa type may be explained by a similar 
development of the mesenteries as in Halcurias or Porponia, though with the 





Fig. 4a. 



Fig. 4b. 




Fig. 4c. 



Fig. 4d. 



FrGS. 4a-d. — Scheme to illustrate the origin of Rugosa, assuming that the development of the mesenteries to begin with 
has proceeded as in the Endocoelactidse. The black tongues are endoccelic septa, the shaded tongues exocoelie septa. 



difference that the development of new mesenteries is continued in the endocoels, and 
that in each quadrant a suppression of mesenteries on the dorsal side takes place in each 



68 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

order from and with the third, a suppression which probably stands in connection with 
the animal's mode of life. 

In this hypothetical explanation of the arrangement of the Rugosa septa, I have 
mainly intended to direct attention of palaeontologists to the fact that the Rugosa type 
may be explained in various ways ; moreover, a closer examination of Rugosa has 
shown that the development is different in different genera, that some of them retain a 
bilateral and others a more radial arrangement ; the development of septa in this 
group, however, requires further examination. 

The hypothesis put forward by me with regard to the origin of Rugosa seems to me 
to speak for itself. The presence of the 4 growth-zones after the development of the 
first 12 mesenteries in Rugosa may be fully explained by supposing a development of 
certain mesenteries like those in the Endocoelactidse. The enlargement of the 4 
primary lateral endoccels has led to developmental zones for the new mesenteries being 
removed to these areas instead of to the 6 primary exoccels. The origin of a 4 (8)- 
rayed type in certain Rugosa may be explained in this way. In any case, I consider 
the above explanation as good as, if not better than, that put forward by Duerden, to 
the effect that Rugosa must stand in a certain relation to the Zoantharia (Zoanthidse). 
In consequence of this view, he also considers the latter group as very old, a view, 
however, I have some difficulty in accepting, as the Zoanthidse are obviously rich in 
species, and presumably form a group which is still in process of differentiation. See 
also my work in Bronn. 

Finally, it seems convenient further to characterise the family Endocoelactidse with 
the genera Halcurias. Porponia, and Synhalcurias. 

Family Endocoelactidse. 

Athenaria (Protactininse?) with thick, sometimes cartilaginous body-wall, without 
sphincter and fossa, with spirocysts in the ectoderm of the body-wall and cesophagus. 
Arrangement of the mesenteries quite different from that of the normal Actiniaria type, 
owing to the development of the second and third order of mesenteries in the endoccels. 
In consequence, the arrangement of the tentacles very different from the normal type 
(among others, 10 tentacles of the first order immediately border on those of the second 
order). Sexual organs present on all the stronger mesenteries from and with those of 
the first order, including directive mesenteries. 

Genus Halcurias M'Murrich = Endoccelactis Carlgren. 

Endocoelactidse with ca. 68 mesenteries, 36 of which are perfect. Four cycles of 
mesenteries. The mesenteries of the fourth order regularly arranged on each side of 
those of the third order, the mesenteries of the same pair equally developed. The 
perfect mesenteries arranged as follows : 20 (6 + 4 pairs) +16 (8 pairs), of which the 
first 20 strong, extending over the whole length of the body ; the others only developed 
in the distal part, and weak. The body cylindrical. The tentacles not bridge-like 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 69 

thickened on the outer side. On the first 20 mesenteries the pennons of the longi- 
tudinal muscles well developed. One oesophageal groove. 

H. pilatus M'Murrich. 

H. carlgreni * M'Murrich [Endocoelactis sp. Carlg. ). 

Genus Porponia R. Hertwig. 

Endocoelactidse with (54? to) ca. 68 mesenteries, of which 44 perfect. Five cycles 
of mesenteries. The mesenteries of the fourth and fifth order are regularly arranged, 
but show unequal development, so that the mesenteries of the fourth order on the one 
side of the mesenteries of the third order consist of a perfect and an imperfect 
mesentery, on the other side only of an imperfect mesentery ; but the mesenteries 
of the fifth order are not paired, and only developed between the mesenteries of the 
first order and the perfect mesenteries of the fourth order (as in Actinostola). The 
arrangement of the perfect mesenteries is 20 (6 + 4 pairs) + 16 (8 pairs) +8 (these 
form pairs with imperfect mesenteries). The body goblet-like, sometimes cylindrical. 
The tentacles on the outer side bridge-like and greatly thickened. The pennons of the 
longitudinal musculature on the mesenteries hardly indicated. Two oesophageal grooves. 

P. elongata R. Hertwig. 
P. robusta R. Hertwig. 
P. antarctica Carlgren. 

Genus Synhalcurias Carlgren. 

Endoccelactidse with considerably more than 68 mesenteries (ca. 100), all of which 
are perfect, arranged in pairs, and frequently agreeing in the size and distribution of 
the sexual organs. The irregular arrangement of the mesenteries probably due to the 
development of the mesenteries of the second and third order in the endocoels. Origin 
of the mesenteries of the fourth order and the following (?). The body cylindrical. The 
tentacles are not thickened on the outer side. The longitudinal muscles of the mesen- 
teries weak, not forming pennons, and almost equally developed on all mesenteries. 
One oesophageal groove (2 ?). 

S. elegans ( WassilierT). 



In a coming work I intend to give a description of the other Actiniaria, ca. 20 in 
number, which have been collected by the Scotia Expedition. 

* As further characterisation of this species, I may give the following information about the nematocysts : — 
Spiroeysts occur in quantities, especially in the tentacles, but are also common in the body-wall, the ectoderm of the 
oesophagus and in the filaments. They are of greatly varying sizes, generally as large as the corresponding thick- 
walled nematocyst capsules ; but smaller as well as still larger ones occur, the latter especially in the tentacles, where 
they reach a length of up to 43 n, breadth 7 p. In the body-wall the thick-walled nematocysts reach a length of 
22-26 fi, in the tentacles 26-34 n, and in the filament and oesophagus ca. 26-29 n. In the latter places are also found 
nematocysts with distinct basal part to the spiral thread, of almost the same length as the preceding, but broader at 
the basal end. The thick-walled nematocysts are most numerous in the tentacles. 



70 PROFESSOR OSKAR CARLGREN ON THE GENUS PORPONIA 

APPENDIX. 

Now that I have had occasion to examine two highly retracted and badly preserved 
specimens of the type of Actinernus, A. nobilis Verr. (place of discovery 43° 18' N., 
60° 24' W., Gloucester Fisheries, 1879, U.S. Fish. Com.), as far as I can see from the 
bad material, the tentacles of the first and second order are arranged as in Porponia. 
There are also spirocysts in the ectoderm of the body-wall. Therefore I think that 
Actinernus nobilis (but not Polysiphonia tuberosa, and probably not A. plebeius — 1 
have not seen this latter species) must be placed in the Endocoelactidse. Whether 
Porponia and Actinernus are synonyms I cannot say for the present, but it is not 
impossible. 

February 4, 1914. 



BIBLIOGRAPHY. 



Carlgrbn, 0., " Studien iiber nordische Actinien," K. Svenska Vet.-Akad. Handl., Bd. xxv., No. 10, 1893. 

"Zur Ketmtnis der Minyaden," O/vers. K. Vet.-Akad. Fork., Stockholm, 1894. 

" Zur Mesenterienentwicklung der Aktinien," O/vers. K. Vet.-Akad. Fork., Stockholm, 1897. 

" Ostafrikanische Actinien," Mitt. Nat. Mus. Hamburg, xvii., 1900. 

" Anthozoa," Bronn's Klassen und Ordnunyen, L. 1-6, 1903-1908. 

" Studien iiber Regenerations- und Regulationserscheinungen, 1, 2," K. Svenska Vet.-Akad. Handl., 

Bd. xxxvii., 1904, Bd. xliii. 1909. 
Duerdkn, J. E., "Relationships of the Rugosa (Tetracoralla) to the living Zoanthese," John H opkins Univers. 

Circul., Jan. 1902. 

"The Antiquity of the Zoanthid Actinians," Sixth Ann. Report Mich. Acad. Sc, 1904. 

Hertwig, R., " Die Actinien der Challengerexpedition," Jena, 1892. 

M'Murrich, J. P., " Report on the Actiniae collected by the United States Fish. Com. Steamer Albatross 

1887-1888," Prod. U. Stat. Nat. Mus., xvi., 1893. 

" Report on the Actiniaria collected by the Bahama Expedition of the State University of Iowa, 1893," 

Ball. Lab. Nat. Hist. Univ. Iowa, 1898. 

" Contributions on the Morphology of the Actinozoa, 6. Halcurias pilatus and Endoccelactis," Biol. Bull., 

ii., No. 4, 1901. 
Pax, F., "Studien au westindischen" Actinien," Zool. Jahrb., Suppl. 2, H. 2, 1910. 
Wassilieff, A., " Japanische Actinien," Abh. Bayr. Acad. Wiss. Mat. Nat., Classe, 1908, 1 Supplb. 



EXPLANATION OF PLATE IV. 

bac, thickenings of the basal part of the ectoderm cells? 

cm., circular muscles. 

ct., ciliated tract of the filament. 

dp., directive plane. 

ec, ectoderm. 

en., entoderm. 

lm., longitudinal muscles. 

m. mesoglcea. 

inf., fibrillous folds of the mesoglcea. 



AND RELATED GENERA, SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 71 



mp., 


muscles' pennon. 


nd., 


enido-glandular tract of the filament. 


pbm., 


parieto-basilar muscles. 


pm., 


parietal part of the longitudinal muscles. 


psb,, 


pseudo-basilar muscles. 


rm., 


radiated muscles of the disc. 


sp., 


spirocysts. 


t, 


tentacles. 



Figs. 1-4, Synhalcurias elegans ; 5-9, Porponia antarctica; 10. Halcurias carlgreni. 

Fig. 1. Transverse section through the body-wall with a part of two mesenteries. £.* 

Fig. 2. Vertical section through the body-wall. Only a part of the mesoglcea is designed. J with out- 
drawn tube. 

Fig. 3. Transverse section of the mesoglcea and muscles of the disc. §. 

Fig. 4. The basal part of a mesenterium with the longitudinal muscles and the pseudo-basilar muscles. 
Schematic. 

Fig. 5. Cross-section through one tentacle above the middle. ^. 

Fig. 6. Cross-section through the same on the basis §. 

Fig. 7. Cross-section through a not-directive mesenterium. The same section as in fig. 8. The whole 
breadth of the mesoglcea is not drawn. §• 

Fig. 8. Cross-section through a part of a directive mesenterium and the body-wall in the region of th e 
aboral end of the stomatodseum. £. 

Fig. 9. Transverse section of the ciliated tract region of the filament, -jy. 

Fig. 10. Section through the upper part of the body to show the arrangement of the mesenteries. Twice 
magnified. 

* Magnifications refer to Reichert's system, " Austria." Figures drawn in the level of the micro- 
scope's foot. 



Trans. Roy. Soc. Edin r 




Carlgren : "Scotia" Genus Pokponia. 






Vol. L.— Plate IV. 



,-fiiS§i 



• • ,■■■4-. .'< : ' ' ,. ■' 'L 



,),:•' t>i ■> 



' ); 



.1 . 

■- : ' : '.-.■..•'■'■. 
V;AV; } >' ■". r -' ' - ■"' ' 

. - - \ i ■ ■ 



- m 








' ■ > < 



\ '• > I 




Oskar Carlgren. del. 



M'F.irlanei Elskiue, I.ith.. Edin. 



( 73 ) 



V. — On the Fossil Flora of the Staffordshire Coal Fields. By R. Kidston, 

LL.D., F.R.S. 

(Read November 17, 1913. MS. received November 24, 1913. Issued separately March 12, 1914.) 

[Plates V.-XVL] 



PART III. 

THE FOSSIL FLORA OF THE WESTPHAL1AN SERIES OF THE 
SOUTH STAFFORDSHIRE COAL FIELD. 

Introduction. 

The first of this series of papers, that on the "Fossil Plants collected during the 
sinking of the shaft of the Hamstead Colliery, Great Barr, near Birmingham," was 
published in 1888,* and the second, dealing with the "Fossil Flora of the Coal Field 
of the Potteries," in 1891.t 

Since that date the North Staffordshire Coal Field has been re-surveyed by members 
of the Geological Survey ,} and the result of this re-examination of the geology of the 
Potteries Coal Field was the classification of the strata into several well-defined 
groups. These groups are given in the following table, and opposite them, in the 
third column, the terms employed in my paper on the " Fossil Flora of the Potteries 
Coal Field " :— 



Thickness. 


Potteries Coal Field. 


Earlier Divisions used in Paper 

on " Fossil Flora of the Coal 

Field of the Potteries." 


Over 700 feet. 
300-350 „ 
800-1100 „ 

300-450 „ 


Keele Group. 

Newcastle-under-Lyme Group. 
Etruria Marl Group. 

Black Band Group. 
Middle Coal Measures. 

Lower Coal Measures and 
Millstone Grit. 


> Upper Coal Measures. 

i Middle Coal Measures (from 
< Brassey Mine Ironstone to 
( Ash Coal). 

Lower Coal Measures (from 
below Ash Coal to top of 
Millstone Grit). 



Subsequent investigations of the fossil plants of the rocks which I there classified 
as Upper Coal Measures showed me that they must be removed from that series — with 
the exception of the Keele Group, which I think is best regarded as the base of the 

* Trans. Roy. Soc. Edin., vol. xxxv. p. 317. + Trans. Roy. Soc. Edin., vol. xxxvi. p. 63. 

I Memoirs of the Geological Survey, England and Wales : The Geology of the North Staffordshire Goal Fields, by 
Walcot Gibson, B.Sc, F.G.S., with contributions by G. Barrow, F.G.S., and C. B. Wedd, B.A., F.G.S. And 
a Palceontological Account, with list of Fossils, by John Ward, F.G.S. , 1905. 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 10 



74 DR ROBERT KIDSTON ON THE 

Upper Coal Measures — and be treated as a distinct passage series, and for them I 
proposed the name of Transition Series* 

The term "Transition Series" was, however, rather indefinite, and also had been 
previously used for rocks of Lower Carboniferous age in Germany, though now seldom 
or never applied to them in recent works. As my work proceeded, I found that this 
" Transition Series " occupied a much more important place in British Upper Car- 
boniferous Rocks than I originally suspected ; so, to prevent any confusion by the 
use of the term " Transition Series," the term Staffordian Series was adopted for the 
inclusion of the Newcastle-under-Lyme, Etruria Marl Group, and the Black Band 
Group as this series is perhaps better developed in that county than in any other 
district of Britain. The Keele Group, being regarded as the base of the Upper Coal 
Measures, is excluded from the Staffordian Series and placed in the group to which 
the name of Radstockian Series was given, t That the Radstockian Series is part of 
the Upper Coal Measures or Stephanian of the Continent is proved by the fossil 
flora of the Radstock Series, which has given origin to the term. 

It is not my intention to enter here largely into the geology of the Staffordshire 
Coal Fields ; but, owing to the period which has elapsed since Parts I. and II. of those 
papers dealing with their' fossil flora were published, it has been necessary to give 
a short account of the change of classification which it has been deemed advisable to 
make, and to show the relationship of the older to the newer terms now in use. 

When writing the first paper of the series, that on the " Fossil Plants found in the 
Hamstead Boring," the following divisions were employed : — 

Upper Coal Measures . . . From the surface to a depth of 1353 feet. 

Middle Coal Measures . . . The remaining portion of section from 

which I saw fossils, down to Heathen 
Coal at depth of 1893 feet. 

To these divisions I must shortly refer. In a discussion on Dr Walcot Gibson's 
paper on " A Boring for Coal at Claverley, near Bridgenorth, and its bearing on the 
extension westwards of the South Staffordshire Coal Field," read before the South 
Staffordshire and Warwickshire Institute of Mining Engineers, at the general meeting 
at Birmingham, February 17, 1913, J Mr F. G. Meachem stated in regard to this 
section that he had asked the late Mr C. E. de Rance some twenty-five years ago to 
mark where he thought the "Red Coal Measures" ended, and he drew the line at 
600 feet from the surface, thus cutting off as Permian 600 feet I had referred to the 
Upper Coal Measures. As a matter of fact, no plants that could be identified were 
found till a depth of 729 feet was reached, and the upper portion of the section may be 
referable to the Permian ; but Mr Meachem does not state on what evidence Mr de Rance 
came to this conclusion. When I examined the plants from this section, now twenty-five 
years ago, we were only beginning to feel our way in the classification of these rocks. 

* Proc. Roy. Phys. Soc. Edin., vol. xii. p. 228, 1894. 

t Quart Journ. Geol. Soc, vol. lxi., 1905, pp. 308-321. 

\ Trans, of the Institute of Mining Engineers, vol. xlv. part i. pp. 30-48, 1913. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 75 

Turning now to the South Staffordshire Coal Field, the following table shows the 

formations represented in that area : — # 

Upper Carboniferous : 

Radstockian Series — 

Keele Beds ....... 800 feet or more. 

Staffordian Series — 

Newcastle Beds . . . . . .500 feet. 

Ktruria Marls . . . . . . 600 to 1000 feet. 

Black Band Group ...... Absent. 

Westphalian Series (Middle Coal Measures) . . . 500 to 2000 feet. 

Lanarkian Series (Lower Coal Measures and Millstone Grit) . Absent. 

Lower Carboniferous ...... Absent. 

Devonian t . . . . . . . . . Present. 

Silurian ........ Present. 

With the plants of the Radstockian Series and Staffordian Series in South 
Staffordshire I do not treat here. At present these rocks are being examined by 
members of the Geological Survey, so it is unnecessary for me to refer to them further 
than to point out (which has already been done in the above table) their relationships 
to the underlying Westphalian Series, with the fossil plants of which the present 
memoir only deals. 

The exposed Coal Measures of South Staffordshire form a tract extending from 
the Clee Hills in the south to Brereton in the north, and from Wolverhampton in the 
west to Walsall in the east, a distance of 21 miles from south to north and about 7 from 
east to west ; but within the last forty years the coal field has been proved to extend 
eastward under the newer rocks, and the Thick coal is now worked at Sandwell and 
Great Barr near Birmingham, and in the last ten years the extension of the coal field 
westwards has been proved at Baggeridge Colliery, Himley. 

With the object of showing the distribution of the fossil plants in the Westphalian 
Series of the South Staffordshire Coal Field, and the relationship of the beds to each 
other from which the plants were derived, a table of the chief strata and coal seams, 
which is taken from the late Mr J. Beete Jukes' memoir on The South Staffordshire 
Coal Field, is inserted here : — \ 

General Section of the Central and Southern Part of the Coal Field. 

/la. The Halesowen \ 
1. § Beds above the Upper Sulphur coal<J Jft ^^^ ^^l > ■ • . 600 to 800 feet. 

' Measure Clays. ) 

* Gibson, Geol. of Coal and Goal Mining, Arnold's Geological Series, 1908, pp. 180-184. Vernon, Quart. 
Journ. Geol. Soc, vol. Ixviii., 1912, p. 613. Kay, Quart. Journ. Geol. Soc, vol. lxix. p. 433, 1913. 

tW. Wickham King, "The Uppermost Silurian and Old Red Sandstone of South Staffordshire," Geol. Mag., 
dec. v., vol. ix. pp. 437-443, 484-491, 1912. 

X Memoir of the Geological Survey of Great Britain and of the Museum of Practical Geology : The South 
Staffordshire Coal Field, by J. Beete Jukes, M.A. Camb., F.R.S., etc., 2nd ed., 1859, p. 20. 

§ These rocks are now included in the Staffordian Series, and include the Keele Group, the Newcastle-under- 
Lyme Group, and the Etruria Marls. 



76 



DR ROBERT KIDSTON ON THE 



2. 

3. 

4. 

5. 

6. 

7. 

8. 

9. 
10. 
11. 



2 < 






13. 

14. 

15. 
16. 
17. 
18. 



19. 

20. 
21. 

22. 
23. 

24. 
25. 
26. 

27. 
28. 
29. 



Upper Sulphur coal 
Intermediate measures 
Little, or Two-foot coal 
Intermediate measures 
(I.) Brooch coal . 

(I. 1) Brooch binds, ironstone measures 
Herring coal (not known north of Dudley) 
(I. 2) Pins and Pennyearth ironstone measures 
Intermediate measures containing the sandstone known as Thick coal rock 
(1. 3) Broad earth, Catch earth, and Batt, containing the Ten-foot, and Backstone 
ironstones in the Pensnett district 
(II.) Roofs coal, or Top floor 
(III.) Top slipper, or Spires, or Spin coal. 

These two form the Flying reed when separated from coals below 
(IV.) Jays, or White coal 
(V.) Lambs, or Floors, or Fine Floors coal. 

These two are often either mentioned together under the name of 
"White coal," or else the lower one is absent 
(VI.) Tow (tough), or Heath coal 
(VII.) Benches coal (this bed is but rarely mentioned) 
(VIII.) Brassils, or Corns coal 
(IX.) Foot coal, or Bottom Slipper, or Fine coal 
(X.) John coal, or Slips, or Veins coal 
(XL) Stone, or Long coal 

(XII.) Patchells coal (sometimes absent or not mentioned) 
(XIII.) Sawyer, or Springs coal 
(XIV.) Slipper coal 
(XV.) Bottom Benches, or Omfray (Humfray), or Red, or Km (Kick?), or 
Holers coal 
(I. 4) Pouncill batt, Blactery and Whitery, containing the Grains ironstone, and 

sometimes the Whitery ironstone 
(I. 5) Gubbin ironstone measures, sometimes called the Little, or Top, or Thick 

coal gubbin, sometimes the Black ironstone 
Table batt and intermediate measures ...... 

(XVI.) Heathen coal .... . . . 

Intermediate measures (sometimes wanting) ..... 

(XVII.) Rubble, or Lower Heathen coal, sometimes, when the measures above 

are wanting, forming the bottom part of the Heathen coal, sometimes 

itself wanting, when the measures above and below seem to be both 

present 

(I. 6) Intermediate measures containing, at Bentley, the ironstones known as the 

Lambstone and Brownstone 
(I. 7) New Mine, or White ironstone ...... 

(I. 8) Measures containing the Pennystone ironstone, called also Bluestone or 

Cakes 
(XIX.) Sulphur or Stinking coal 
Intermediate measures ..... 

(XX.) New Mine coal ..... 

(I. 9) Measures containing Fireclay Balls ironstone (occasionally) 

(XXI.) Fireclay coal (and partings) 

Intermediate measures ..... 

(I. 10) Getting Rock ironstone (occasionally) 
(I. 11) Poor Robin ironstone measures 



about 1 foot. 
140 feet. 
2 feet. 

from 2 to 48 feet, 
about 4 feet, 
from 7 to 20 feet, 
about 1 foot, 
from 7 to 30 feet. 
„ 38 to 157 feet. 

• ,, 6 to 14 feet. 



\ about 30 feet. 



• from 2 to 8 feet. 

• „ 2 to 8 „ 

„ 2 to 28 „ 
about 3 feet, 

from to 43 feet. 

v „ 2 to 4 „ 

• „ 10 to 33 feet. 
„ 2 to 10 

> „ 10 to 25 

„ 2 to 9 

„ 3 to 99 

,, 2 to 11 

„ 2 to 40 

„ 1 to 14 

„ 2 to 10 

„ 4 to 5 

„ 3 to 5 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 



77 





from to 19 feet 






, 2 to 9 „ 






, 3 to 12 „ 






, 5 to 30 „ 


■ Bottom Gubbin 




, 3 to 10 „ 
, 18 to 50 ,. 
, 2 to 4 „ 
, 16 to 50 „ 
, 2 to 9 „ 
, 10 to 14 „ 
, 4 to 7 „ 
, 6 to 15 „ 
, 2 to 3 „ 


s ironstone 


about 50 feet. 



30. Intermediate measures, sometimes wanting 

31. (I. 12) Rough Hills White ironstone (occasionally) 

32. (XXII.) Bottom coal ..... 

33. Intermediate measures ..... 

34. (I. 13) Gubbin and Balls ironstone, sometimes called the Great or 

35. Intermediate measures ..... 

36. (XXIII.) Singing, or Mealy Grey coal (occasionally) 

37. Intermediate measures ..... 

38. (I. 14) Blue Flats ironstone .... 

39. Intermediate measures ..... 

40. (I. 15) Silver Threads ironstone .... 

41. Intermediate measures ..... 

42. (I. 16) Diamonds ironstone .... 

43. Lowest measures, -maximum thickness known below the Diamonds ironstone 

It must be mentioned, as pointed out by its compiler, that the whole section does 
not occur in any one point of the coal field, some parts of the section being absent 
in one area and present in another, and at no one point are all these beds to be found. 
The total estimated thickness of the Westphalian Series in the northern division 
of the South Staffordshire Coal Field is about 2000 feet, and in the southern portion 
between 500 and 600 feet. # This is brought about by the splitting up of the 
Thick coal, which attains a thickness of 30 feet in the southern area, into eight or nine 
seams, which, as traced into the northern division, become more and more separated 
by an increasing thickness of the intermediate measures. 

The whole extent of the coal field as at present known is about 149 square miles.t 

Very little has been published on the fossil plants of the Westphalian Series of 
South Staffordshire, the only notes known to me being that by Hooker in his paper, 
'" On the Vegetation of the Carboniferous Period, as compared with that of the Present 
Day" ; J a short paper by Mr B. Smith, § in which he figures a Lepidodendron stem which 
may possibly be the Lepidodendron distans Lesqx. ; and a " Note on the Occurrence of 
Whittleseya elegans Newb. in Britain," by Mr H. Hamshaw Thomas. || In addition 
to these are the three papers by myself : that on the fossil plants collected during the 
sinking of the shaft of the Hamstead Colliery, Great Barr, near Birmingham, to which 
reference has already been made ; the description of the fructification of Neuropteris 
heterophylla Brongt.;1I and the description of the microsporangia of Sphenopteris 
Honinghausi Brongt.** 

From the northern portion of the coal field the plant records have not been so 
fully worked out as in the southern area. 

Since beginning the study of the fossil plants of the South Staffordshire Coal 

* Gibson, The Geology of Goal and Goal Mining, Arnold's Geological Series, 1908, p. 184. 

t Gibson, loc. cit., p. 149. 

J Mem. Geol. Survey of Great Britain, vol. ii. part ii. p. 387, 1848. 

§ Geological Mag., dec. v., vol. ii. p. 208, 1905. 

|| Palaeobotanische Zeitschrift, vol. i. part i., Nov. 1912, p. 46 ; text-figs. 1, 2. 

IT Trans. Roy. Soc. London, series B, vol. cxcvii. pp. 1-5, pi., 1904. 

** Trans. Roy. Soc. London, series B, vol. cxcviii. pp. 413-445, pis. xxv.-xxviii., 1906. 



78 DE ROBERT KIDSTON ON THE 

Field, thirty years ago, some to whom I am much indebted for kind help have passed 
away ; and I would specially wish to acknowledge the assistance I received from the 
late Messrs Henry Johnson, sen., F.G.S., Henry Johnson, jun., and Charles Beale, 
all of Dudley ; and from Mr John Ward, F.G.S., of Longton. 

To Sir Charles Holckoft, now of Kingswinford ; Mr D. Rodgers ; Mr Henry 
Insley, Great Barr ; Mr W. J. Davis, Bilston ; Mr T. G. Latham, Dudley ; 
Mr W. Madeley, Dudley; Mr W. Egginton, Sedgley ; Dr Wheelton Hind, F.G.S., 
and Mr J. T. Stobbs, F.G.S., both of Stoke-upon-Trent, my thanks are also due for 
much help ; but above all am I indebted to Mr H. W. Hughes, F.G.S., Dudley, who 
during the whole of the long period in which I have studied the fossil plants of the 
South Staffordshire Coal Field has availed himself of every opportunity for collecting 
them, and through whose labours I am able to record here many of the species 
included in this memoir. To him also we are indebted for the discovery of the seed of 
Neuropteris heterophylla Brongt., and the microsporangia of Sphenopteris (Crossotheca) 
Honinghausi Brongt. 

LIST OF SPECIES. 
FERNS AND FERN-LIKE PLANTS. 

Sphenopteris Brongniart. 
Sphenopteris obtusiloba Brongt. 

1829. Sphenopteris obtusiloba, Brongt., Hist. d. veget. foss., p. 204, pi. liii. fig. 2. 

1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 65, pi. iii. figs. 1-4; pi. iv. 

fig. 1 ; pi. v. figs. 1, 2. 

1911. „ ,, Kidston, Mem. Musee roy. d'hist. not. de Belgique, vol. iv. p. 9. 

1833. Sphenopteris irregularis, Sternb., Essai flore monde prim., vol. ii., fasc. v., vi., p. 63, pi. xvii. 

fig. 4. 
1885. Diplothmema Schumanni, Stur, Die Fame : Carbon-Flora d. Schatz. Schichten, p. 352, pi. lxv. 

fig. 2. 
1885. Diplothmema Schlotheimii, Stur (non Brongt.), ibid., p. 336, pi. xxv. fig. 4. 

1912. Sphenopteris Schumanni, Gothan, Verhandl. d. naturhist. Vereins d. preuss. Rheinl. u. Wesi- 

phalens, 69. Jahrg., p. 240, pi. iii. figs. 1, 2. 

Remarks.- — Some difference of opinion exists amongst botanists as to the specific 
position of Diplothmema Schumanni Stur ; some regard it as a distinct species, and 
others as synonymous with Sphenopteris obtusiloba Brongt., and this latter view is 
that accepted here. 

Dr Gothan has recently figured specimens of Sphenopteris Schumanni Stur,* 
and in a tabular form points out what he regards as the characters which separate it 
from Sphenopteris obtusiloba Brongt. He also gives a column showing wherein 
Sphenopteris trifoliolata Artis differs from these two species ; but the plant he has 

* Loc. cit. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 79 

accepted as Sphenopteris trifoliolata is not the Sphenopteris trifoliolata of Artis, but, 
I believe, only a form of Sphenopteris obtusiloba Brongt. The true Sphenopteris 
trifoliolata Artis does not seem to have been found outside of England. 

That portion of his table which refers to the plants under discussion is given here :— 



Sphenopteris Schumanni. 
Pinnules convex, round. 
Pinnule upper surface smooth. 

Nervation standing out in prominent relief. 

Rachis with broad furrow. 
Rachis smooth. 



Sphenopteris obtusiloba. 
Pinnules flat, round. 
Pinnule upper surface covered with close 

radiating striations. 
Nervation scarcely visible, generally obscured 

by the radiating striations. 
Rachis flat without furrow. 
Rachis with numerous transverse bars. 



With the object of studying this question, through the kind offices of Dr Gothan 
I received from Dr A. Franke, Dortmund, examples of Sphenopteris Schumanni from 
Zeche Gluckauf Tiefbau, near Dortmund, the same locality as that from which the 
specimens figured by Dr Gothan in his pi. iii. figs. 1, 2 were derived. I am also 
indebted to Dr Beyschlag, Director of the Konigliche Geologische Landesanstalt, 
Vienna, for kindly sending me for examination a specimen of Sphenopteris Schumanni 
from the collection under his charge. I cannot therefore be mistaken as to the plant 
identified by Stub, and Gothan as Sphenopteris Schumanni, and in fact my own 
collection, under the name of Sphenopteris obtusiloba Brongt., contains specimens that 
would be named Sphenopteris Schumanni by those who uphold this as a species. I find 
myself, however, unable to regard Sphenopteris Schumanni as specifically distinct, for 
among the specimens I have examined I find that the supposed distinctive characters are 
not constant, and are apparently due in part to causes attending their fossilisation. 

If one now considers these supposed specific differences, first, as to the pinnules 
being convex in Sphenopteris Schumanni and flat in Sphenopteris obtusiloba, it will be 
found, I think, that this depends entirely on the amount of pressure to which the 
fossils have been subjected. On the specimen of Sphenopteris Schumanni from Buhen 
Grube near Neurode, 7 Flotz, kindly lent me by Dr Beyschlag for examination, at one 
side of the slab a few of the pinnules are more or less convex, while on the other side some 
of the pinnules are quite flat and the rachis is perfectly smooth without any trace of the 
" broad furrow," this condition of the specimen being evidently the result of pressure. 

On the example of Sphenopteris obtusiloba figured in the Trans. Roy. Soc. Edin., 
vol. xxxvii. pi. i. fig. 1, # the pinnules are flat and the nervation well shown on account 
of the striated epidermal surface having been removed from all the pinnules except 
at one or two points. The rachis here is also smooth and shows no trace of the trans- 
verse bands which are one of the distinctive characters of Sphenopteris obtusiloba. The 
surface striation of the pinnules is caused by the arrangement of the epidermal cells. 

It is seen, then, that the characters said to be available for the separation of 
Sphenopteris Schumanni Stur from Sphenopteris obtusiloba Brongt. are most 

* "On the Fossil Plants of the Kilmarnock, Galston, and Kilwinning Goal Field," p. 307, pis. i.-iv. 



80 DR ROBERT KIDSTON ON THE 

inconstant in both " species," and even at the best they seem to be the result of 
secondary causes, and not dependent on original structural differences. 

Sphenopteris Schumanni is therefore here united with Sphenopteris obtusiloba 
Brongt. 

Horizons and Localities. — 

Roof of Brooch Coal: Himley and Tividale. 

Ten-foot Ironstone Measures : Clayscroft Open Work, Coseley, near Dudley. 

Roof of Thick Coal: Bradley Colliery, Bilston. 

Roof of New Mine Coal : Doulton's Clay Pit, Netherton ; Mount Pleasant, 

Brierley Hill. 
Roof of Bottom Coal : Coseley. 

Sphenopteris dilatata L. & H. 

PL V. figs. 1 and la. 

1832. Sphenopteris dilatata, L. & H., Fossil Flora, vol. i. pi. xlvii.* 

1861. Sphenopteris Honinghausi, Salter (non Brongt.), Mem. Geol. Surv. of Great Britain: Iron Ores of 
Great Britain, part iii.; Iron Ores of South Wales, p. 232, fig. 2. 

Remarks. — It is extremely difficult, probably impossible, to ascertain the true 
characters of Sphenopteris dilatata as figured and described by Lindley and Hutton. 
The type appears to be lost, and, notwithstanding what is stated in the description 
in regard to the nervation, there seems to be no doubt the plant is a typical 
Sphenopteris, and the enlargement which accompanies their figure bears out this view. 
According to the enlarged figure, the nervation is very close, but I strongly suspect 
that the artist is responsible for this, for unfortunately, when one is enabled to 
compare the figures of the Fossil Flora with the types, the former are often found 
to be most inaccurate ; hence I am not inclined to place too much dependence on the 
accuracy of this figure. 

On Plate V. at fig. 1 is given a small specimen of the plant I believe to be the 
Sphenopteris dilatata L. & H. It agrees well with the general character of the plant. 
The pinnules are very convex and their distal margin is suddenly bent down, which 
gives them a much more truncate appearance than they really possess. This is shown 
somewhat in the enlargement given by Lindley and Hutton. If the pinnules were 
spread out, their form would be, I believe, that of Sphenopteris obtusiloba Brongt. 
They have very wide foot-stalks, into which only one vein appears to enter from the 
rachis, but perhaps they are not wider than seen on the pinnules of the uppermost 
pinnae of Sphenopteris obtusiloba. Several different localities have yielded specimens 
of the plant I refer to Sphenopteris dilatata, but they are all upper portions of the 
frond or of the large primary pinnae, and I am now inclined to think that the species 

* The Sphenopteris dilatata Lesquereux in Owen, 2nd Rept. Geol. Reconnoissance of Arkansas, p. 310, pi. iii. fig. 3, 
1860, is not Lindley and Hutton's species of that name. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 81 

is founded on an incompletely developed condition of Sphenopteris obtvsiloba, though 
one hesitates to affirm that this is the true interpretation of the species. 

I originally referred Sphenopteris dilatata to Sphenopteris trifoliolata Artis, sp., 
but in this I now feel certain I was in error.* 

The fossil figured as Sphenopteris Honinghausi by Salter f should, I think, be 
referred to Sphenopteris dilatata, whatever position may be eventually assigned to 
that plant. 

My thanks are due to Dr A. Smith Woodward, F.R.S., for permission to figure 
the specimen shown on the Plate, which is preserved in the Geological Department 
of the British Museum, No. V., 1377. 

Horizon and Locality. — Ten-foot Ironstone Measures {—Roof of Thick Coal): 
Coseley, near Dudley. 

Sphenopteris trifoliolata Artis, sp. 

1825. Filicites trifoliolatus, Artis, Antedil. Phyt., p. 11, pi. xi. 

1828. Sphenopteris trifoliolata, Brongt., Prodrome, p. 50. 

1836. Cheilanthites trifoliolatus, Gopp., Sys. fil. foss., p. 245. {Refs. in part.) 

1911. Eusphenia trifoliolata, Anon., Catal. Coll. foss. conserves an Mus. liouil. de Lille, p. 26. 

Remarks. — Up to the present, this species does not appear to have been found 
outside of England ; for though several Continental botanists have figured a 
Sphenopteris under the name of Sphenopteris trifoliolata Artis, sp., none of the 
figures they give are referable to Artis's plant. \ 
Horizons and Localities. — 

Blue Measures, six feel above Brooch Coal: Hamstead Colliery, Great Barr, 

near Birmingham. Collected by Mr H. Insley. 
Roof of Brooch Coal: "Neighbourhood of Dudley " (? Himley). 

Sphenopteris Sauveuri Crepin. 

1880. Sphenopteris Sauveuri, Crepin, Soc. roy. hot. de Belgique, vol. xix. part ii. p. 17. 

1886. „ „ Zeiller, Flore foss. bassin houil. d. Valen., p. 79, pi. ix. fig. 6. 

1903. ,, „ Potonie, Abbild. u. Beschreib. Pflanzen-Reste, Lief. i. No. 4, figs. 1-3. 

1829. Sphenopteris Schlotheimii, Brongt. {non Sternb.), Hist. d. veget. foss., vol. i. p. 193, pi. Ii. {Excl. 

syn.; fig. imperfectly drawn.) 
1848. Sphenopteris elegans, Sauveur {non Brongt.), Veget. foss. de la Belgique, pi. xviii. fig. 3. 
1885. Diplothmema Schlotheimii, Stur, Die Fame d. Schatz. Schichten, p. 336, pi. xx. figs. 1, 2 {non 

pi. xxv. fig. 4). 
1866. Sphenopteris obtusiloba, Andrae {non Brongt.), Vorwelt. Pjtanzen, p. 32, pi. x. figs. 1-4. {Excl. 

refs.) 
1885. Diplothmema Richthofeni, Stur {pars), ibid., p. 343, pi. xxv. figs. 6-7. 

Note. — Sphenopteris Sauveuri is rare in Britain. 

* Catal. Palaeoz. Plants, p. 72, 1886. t hoc. cit. 

% The specimens usually referred to Sphenopteris trifoliolata are, I believe, forms of Sphenopteris obtusiloba. For 
references see Kidston, Mem. Muse'e roy. d'hist. nat. de Belgique, vol. iv. p. 10, 1911. 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 11 



82 DR ROBERT KIDSTON ON THE 

Horizons and Localities. — 

Blue Measures, six feet above Brooch Coal: Hamstead Colliery, Great Barr, 

near Birmingham. Collected by Mr H. Insley. 
? Roof of Thick Coal : Tipton. Specimen in the Collection of the Geological 

Department, British Museum, No. 52,543. 
Ten foot Ironstone Measures : Clayscroft Openwork, Coseley. 

Sphenopteris Laurenti Andrae. 

1869. Sjyhenopteris Laurenti, Andrae, Vorwelt. Pflan?:en, p. 39, pi. xiii. figs. 1-3. 

1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 85, pi. vi. fig. 3, pi. ix. fig. 4. 

1883. Hapalopteris Laurenti, Stur, Morph. u. Syst. d. Culm- u. Carbon-Fame, p. 32. 

1869. Sphenopteris stipulata, Andrae (non Gutbier), ibid., p. 40, pi. xiii. fig. 4. 

Horizon and Locality. — Roof of Fireclay Coal : Doulton's Clay Pit, Netherton, 
near Dudley. 

Sphenopteris Schillingsi Andrae. 
PI. I. figs. 2 and 2a. 

1866. Sphenopteris Schillingsii, Andrae, Vonoelt. Pflanzen, p. 22, pi. vii. figs. 1, la, lb. 
1899. ,, „ Zeiller, F-tude sur la florefossile du bassin d'Heraclee, p. 5. 

Description. — Frond tripinnate, rachis slender. Primary pinnae ascending, 
alternate, subtriangular, obtuse ; secondary pinnae ascending, touching each other 
laterally or slightly distant, alternate ; the lower narrow-deltoid, obtuse, and bearing 
two to three pairs of pinnules. 

Pinnules ascending, slightly decurrent, sometimes inequilateral, alternate or 
opposite, ovate-rotund, with two to three blunt lobes and obtuse apices, the lower 
ones shortly stalked, the upper sessile ; uppermost pinnules sometimes confluent, 
oblong, slightly lobed, more commonly entire, obtuse ; terminal pinnule rotund-cunate. 

Central vein scarcely stronger than the others, and from a short distance above its 
base gives off lateral dichotomous veinlets. Frequently on each side of the central 
veinlet is a lateral veinlet which unites with the central vein at its extreme base or 
joins directly on to the rachis ; all the lateral veinlets arise at an acute angle. 

Remarks. — Although the specimen I refer to this species is a very fragmentary 
one, the size and form of the pinnules agree so well with the corresponding part of 
Andrae's figure that I have little doubt in referring it to Sphenopteris Schillingsi. 
The nervation also shows in one of the pinnules the peculiar character of a lateral vein 
next the mid-rib running directly into the rachis ; while in another pinnule two veins 
of equal strength seem to unite at the extreme base of the pinnule, where it joins 
on to the rachis. 

The small specimen is seen natural size on Plate V. fig. 2, and enlarged two 
times at fig. 2a. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 83 

This is the only British example of Sphenopteris Schilling si Andrae which has 
come under my notice. 

Horizon and Locality. — Ten-foot Ironstone Measures: Cabbage Hall Pits, 
NethertOD. 

Collected by Mr H. W. Hughes, F.G.S. 

Sphenopteris furcata Brongt. 

1829. Sphenopteris furcata, Brongt., Hist. d. veget.foss., p. 179, pi. xlix. figs. 4-5. 

1885. Diplothmerna furcatum, Stur, Carbon-Flora d. Schatz. Schichten : Die Fame, p. 299, pi. xxviii. 

figs. 2, 3. 
1892. Palmatopteris furcata, Potonie, " Ueber einige Carbon fame," part iii. p. 1, pi. i. ; text-fig. 1, p. 5 
(Jahrb. d. kbnigl. preuss. geol. Landesanstalt fur 1891). 

Horizons and Localities. — 

Roof of Thick Coal: Bradley Colliery, Bilston. 

Ten-foot Ironstone Measures : Clayscroft OpenwcJrk, Coseley, near Dudley. 

Immediately below Bottom Coal : Euiton, near Sedgley. 

Sphenopteris spinosa Gopp. 

1842. Sphenopteris spinosa, Gopp., Genre d. plant, foss., parts iii.-iv. p. 70, pi. xii. 

1886. ,, „ Zeiller, Flore foss. d. bassin houil. d. Valen., p. 135, pi. xv. figs. 1-3 

1880. ,, ,, Crdpin, Bull. Soc. roy. de botan. de Belgique, vol. xix. part ii. p. 13. 
1877. Diplothmerna spinosum, Stur, Culm-Flora, vol. ii. p. 230. 

1885. ,, „ Stur, Carbon-Flora d. Schatz. Schichten: Die Fame, p. 312, pi. xxviii. 

figs. 7, 8. 
1869. Sphenopteris palmata, Sehimper, Traite d. paleont. veget., vol. i. p. 388, pi. xxviii. fig. 1. 
1877. Diplothmerna palmatum, Stur, Culm-Flora, veil. ii. p. 230. 

1885. ,, ,, Stur, Carbon-Flora d. Schatz. Schichten: Die Fame, p. 310, pi. xxvii. 

fig. 3. 
1904. Palmatopteris furcata, Potonie (non Brongt.) (pars), Abbild. u. Beschreib. foss. Pflanzen, Lief. 2, 
No. 21, fig. 4 and plate. 

Remarks. — Only one specimen of this species has been collected by Mr Hknry 
Insley. 

Horizon and Locality. — Blue Measures, six feet above Brooch Coal: Hamstead 
Colliery, Great Barr, near Birmingham. 

Sphenopteris artemisisefolioides Crepin. 

1881. Sphenopteris artemisisefolioides, Crepin, in Mourlon, Geol. de la Belgique, vol. ii. p. 60. 

1886. ., ,. Zeiller, Flore foss. bassin houil. d. Valen., p. 132, pi. xiv. 

figs. 2, 3.' 
1910. „ ,, Renier, Paleont. du terr. houil , pi. lxxii. figs, a, b. 

1848. Sphenopteris artemisixfolia, Sanveur (non Sternb.), Veget.foss. terr. houil. de la Belgique, pi. xx. 

figs. 1, 2 (non fig. 3). 
1876. Eremopteris artemisixfolia, Boulay, Terr, houil. du Nord de la France, pp. 28, 73, pi. i. fig. 6. 

1849. Sphenopteris stricta, Sauveur (non Sternb.), ibid., pi. xix. fig. 1. 



84 DR ROBERT KIDSTON ON THE 

1885. Archnopteris Crepini, Stur, Carbon-Flora d. Schatz. Schichten: Die Fame, pi. xxv. figs, la, lb, 

2, and 3. 
1885. Archxopteris Sauveuri, Stur, ibid., pi. xxxvi. fig. 2. 
1893. Sphenopteris spiniformis, Kidston, Trans. York. Nat. Union, part xviii. p. 103. 

Horizon and Locality. — Roof of New Mine Coal: Clattershall Colliery, Brettell 
Lane. 

Sphenopteris Schatzlariana Stur (emend.). 

1885. Diplothmema Schatzlarense, Stur, Die Fame : Carbon-Flora d. Schatz. Schichten, p. 325, pi. xxix. 

figs. 10, 11, pi. lxiv. fig. 2. 
1913. ,, ,, Gothan, Oberschlesische Steinhohlenftora, i. Theil, p. 75, pi. viii. fig. 5. 

Remarks. — The genus Diplothmema Stur, in which he includes the above species, 
was founded for certain ferns or fern-like plants in which more probably the petiole 
than the rachis of the primary pinnae bifurcated ; the part below the fork was naked, 
while the leafy portion was borne on the resulting forks or arms. 

A.s employed, this genus seems to bring together forms characterised in many cases 
by little or nothing in common except a bifurcation of the petiole, and becomes in fact, 
as generally used, practically another name for Sphenopteris Brongniart, if we add 
to that genus the additional character of the bifurcation of the petiole in certain of 
the species, and the use of the genus infers no additional knowledge of the affinities or 
nature of the plants it embraces. 

I am aware that most paheobotanists have adopted the genus Diplothmema in a 
more or less restricted sense than that originally proposed by Stur, but I have not 
seen my way to follow this course, as it does not appear to be any real advance on the 
old form genus Sphenopteris ; and even as a form genus, above which rank Diplothmema 
does not rise, specimens showing the Diplothmema character are so rarely found 
that as a means of provisionally separating certain forms it is of little practical value. 

But the course I adopt raises a difficulty in the case of Diplothmema Schatzlarense 
Stur, for it cannot be simply removed from Diplothmema and placed in Sphenopteris, 
as there is a Sphenopteris Schatzlarensis Stur, sp., already, the Hapalopteris 
Schatzlarensis Stur (pars). As a means, therefore, of getting over this difficulty I have 
altered the specific name from Schatzlarensis to Schatzlariana. 

Sphenopteris Schatzlariana is rare in Britain, but I have already seen it from the 
Yorkshire and North Staffordshire Coal Fields. 

Horizon and Locality. — Roof of Fireclay Coal : Doulton's Clay Pit, Netherton, 
near Dudley. 

Cf. Sphenopteris Sancti-Felicis Stur, sp. 
PI. X. figs. 8 and 8a. 

1885. Diplothmema Sancti-Felicis, Stur, Die Fame: Carbon-Flora d. Schatz. Schichten, p. 301, 
pi. xxix. fig. 1. 



FOSSIL FLOE A OF THE STAFFORDSHIRE COAL FIELDS. 85 

Remarks. — I refer provisionally to this species the small specimen given on PL X. 
fig. 8, natural size, and enlarged 2 times at fig. 8a. 

In the form of the pinnules and their being divided into sharp-pointed, wide- 
spreading segments, it agrees well with Stur's figure ; but it differs in the pinnules 
having a smooth surface, not longitudinally striate as in the type specimen. A single 
vein is clearly seen to enter each segment of the pinnule in the Sandwell Park fossil, 
while the veins are said to be masked by the surface longitudinal striations in Stur's 
example. Whether these differences are specific or due to their condition of preservation 
can only be determined when more perfect specimens are discovered. 
Horizons and Localities. — 

Blue Measures above Brooch Coal: Jubilee Pit, Sandwell Park, West 

Bromwich. 
Ten-foot Ironstone Measures : Bradley Colliery, Bilston. 
Both collected by Mr H. W. Hughes, F.G.S. 

Sphenopteris (Hymenophyllites) quadridactylites Gutbier. 

1835. Sphenopteris quadridactylites, Gutbier, Abdr. u. Ver. d. Zwick. Schwarzk., p. 36, pi. xi. fig. 5. 
1884. Hymenophyllites quadridactylites, Kidston, Quart. Journ. Geol. Soc, vol. xl. p. 596. (Excl. ref., 

Sphenopteris opposita and Sphenopteris minuta Gutbier). 
1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 100, pi. viii. 

figs. 1-3 ; text-fig. 36, p. 56. 
1838. Sphenopteris tetradactyla, Presl, in Sternb., Vers., ii., fasc. vii.-viii. p. 128. 
1855. Sphenopteris tridactylites, Geinitz (non Brongt.), Vers. d. Steink. in Sachsen, p. 15, pi. xxiii. 

figs. 13-14. 
1883. Hymenophyllites delicatulus, Zeiller (non Sternb.), Ann. d. sc. nat., 6 e seV., vol. xvi. pp. 196 and 

208, pi. x. figs. 22-32. 

Horizon and Locality. — Ten-foot Ironstone Measures : Coseley, near Dudley. 



Sphenopteris deltiformis Kidston, n. sp. 
PI. X. figs. 9 and 9a. 

1899. Sphenopteris (Renaultia) bella, Zeiller (non Stur), " Etude sur la More foss. bassin houil. d. 
Heraclee," Me'm. Soc. geol. d. France: Paleont., Mem. No. 21, p. 15, pi. i. fig. 13. 

Description. — Frond quadripinnate, rachis flexuous. slender, and bearing scattered 
spine-like setae. Primary, secondary, and tertiary pinnse deltoid, touching or over- 
lapping at the margins, of somewhat lax growth ; rachis of tertiary pinnse winged, and 
bearing 3-4 pairs of alternate pinnules. Pinnules ovate, the lower bearing 2 pairs 
of lateral lobes and a terminal one, lobes directed forward and separated by a 
sinus extending to near the mid-rib, but the lobes are united to each other at their 
bases, apices of segments rounded or obtusely pointed ; upper pinnules with fewer lobes 
of similar form ; terminal lobe of pinna simple or trifid. The slightly flexuous central 



86 DR ROBERT KIDSTON ON THE 

vein gives off simple or dichotomously divided veinlets, one fork of which goes to 
each lobe. 

Remarks. — The specimen described here and shown natural size on PI. X. fig. 9 
seems to be identical with that figured by Zeiller under the name of Sphenopteris bella, 
but which appears to be specifically distinct from that species. On our plant the 
rachis bears somewhat distant little thorn-like spines, but these, unless the specimen 
were well preserved, might not be shown on the fossil. Irrespective of this character, 
however, the rachis is flexuous, the pinnae deltoid, and the pinnule lobes more spreading 
than those of Sphenopteris (Hapalopteris) bella Stur. # The plant has also a much 
more lax type of growth. If the enlargement of the pinnules given here twice natural 
size on PL X. fig. 9a be compared with the enlargement of the pinnules of Sphenopteris 
bella Stur, the difference will be clearly seen.f In Sphenopteris bella the pinnules are 
smaller, the lobes more truncate and not so spreading, which gives a much denser 
character of growth to the frond. The pinnae are also narrower, not so deltoid as in 
Sphenopteris deltiformis, and the rachis is straight. 

The specimen was collected by Mr H. W. Hughes, F.G.S. 

Horizon and Locality. — Whitestone : Racecourse Pit, Round Oak. 

Sphenopteris Kilimlii Kidston, n. sp. 
PI. V. fig. 3. 

1899. Sphenopteris (Renaultia?) Aschenborni, Zeiller (non Stur), Mem. Soc. geol. d. France, Mem. 
No. 21, p. 14, pi. i. figs. 15, 15a. 

Description. — Frond quadripinnate, primary pinnae lanceolate ; secondary pinnae 
alternate approximate, lanceolate, rachis straight, finely striated longitudinally ; 
tertiary pinnae alternate approximate, lanceolate or oblong-lanceolate, rachis winged 
and bearing 3-8 pairs of alternate pinnules ; pinnules oval or oblong-oval, placed close 
together but seldom touching, contracted into a broad footstalk, slightly directed 
forward, decurrent ; lowest pinnule on upper side (anadrome) bears 3-6 lateral simple 
lobes, with a blunt or bifid point, basal lobe on distal side of pinnule separated from 
tertiary rachis by a narrow sinus, terminal lobe bifid or simple ; basal pinnule on under 
side (catadrome) of rachis occupying a middle position in angle formed by union of 
secondary and tertiary rachises and not touching either with any of its parts ; middle 
pinnules with fewer simple or bifid segments ; uppermost pinnules simple or bifid ; 
surface of pinnules with a roughened appearance. 

Nervation : central vein straight, giving off alternate simple or dichotomous veinlets, 
one branch of which ends in each tooth or segment. 

Remarks. — The lower tertiary pinnae are more broadly lanceolate than those on the 
upper part of the secondary pinnae. This is seen on Plate V. fig. 3, where the 

* Die Fame: Carbon- Flora Hchatz. Schicht., p. 50, pi. xlii. figs. 1, 2. 

t See Kidston, Mem. Muse'e rorj. d'hist. nat. de Belgique, vol. iv. pi. vi. fig. 1. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 87 

fragment of the secondary pinnae shown at the upper part of the specimen bears more 
broadly lanceolate tertiary pinnae than those seen on the secondary pinnae on the right- 
hand lower portion of the figure. These latter originate from secondary pinnae 
which occupied a higher position on the frond than those of the fragment seen at the 
upper margin of the figure. 

The lower pinnules may bear as many as six lateral lobes, three on each side, and in 
these the pinnule is more elongated and is distinctly narrower in proportion to its 
length than those with fewer lobes. These much-divided pinnules are seen on the 
lower tertiary pinnae at the upper end of the figure. 

Text-fig. 1 (a) shows a pinnule from a short distance above the base of the pinna, 





a 
Text-fig. 1. — Sphenopteris Kilimlii Kidston. Two pinnules enlarged 12 times. 

and 1 (&) one towards the apex. The frond has probably been of considerable size, for on 
one of my specimens (No. 1614) what appears to be part of a rachis is 3 "50 mm. thick. 

The surface of the pinnules when seen under a lens has a distinctly roughened 
appearance. Under higher magnification this is seen to be caused by the large size of 
the epidermal cells. The same roughened appearance is shown by Zeiller in the 
enlarged figure he gives of his specimen.* It has not been attempted to represent this 
character in my enlargements. 

Sphenopteris Kilimlii has a slight similarity to Sphenopteris Schatzlarensis Stur, 
sp. {pars),}' but this latter species is distinguished from it by its more deltoid pinnules, 

* hoc. cit., pi. i. fig. 15a. 

t Stur included two species under his Hapalopteris Schatzlarensis in Die Fame d. Carbon-Flora d. Schatzlarer 
Schichten, p. 58. The genus Boweria is founded on specimens similar to Stur : s pi. xl. fig. 1, which has been removed 
from Sphenopteris Schatzlarensis, so that Stur's specimen given on his pi. xl. fig. 1 can now be distinguished as 
Boweria Schatzlarensis. The original description of Boweria will be found in Kidston, Mem. Muse'e roy. d'hist. nat. de 
Belgique, vol. iv. p. 31, 1911. 



88 DR ROBERT KIDSTON ON THE 

which narrow more upwards, and by its sharp-pointed segments, which are also much 
more seldom bifid. The surface of the pinnules is also smooth, not roughened as in Sphen- 
opteris Kilimlii, though imperfect states of preservation may not show this character. 

It has been pointed out by Behrend * and GoTHANt that the plant named 
Hapalopteris Aschenborni by Stur \ is identical with his Ifapalopteris (Sphenopteris) 
Schatzlarensis in part§ — a conclusion with which 1 agree. 

The plant described here is not the Sphenopteris Aschenborni Stur, sp., which is 
synonymous with Splienopteris Schatzlarensis Stur (pars), but the Sphenopteris 
Aschenborni Zeiller (non Stur). It is therefore necessary to give a new name to this 
species, and I propose to distinguish it as Sphenopteris Kilimlii, from the locality at 
which it was originally discovered. 

My thanks are due to Mr H. W. Hughes, F.G-.S., for specimens of this species, 
which has hitherto only been collected at Kilimli. 

Horizon and Locality. — Blue Measures, six feet above Fireclay Coal: Doul ton's 
Clay Pit, Netherton, near Dudley. 

Sphenopteris Waited Stur, sp. 

1885. Calymmotheca Walteri, Stur, Carbon-Flora der ScJiatzlarer Schichten : Die Fame, p. 263, 

pi. xxxvi. fig. 4. 
1893. Palmatopteris Waited, Potonie, " Ueber einige Carbonfarne," iv. Theil, Jahrb. d. kbnigl. 
preuss. geol. Landesanstalt fur 1892, p. 8, pi. iii. 

Remarks. — Apparently rare, but found at Hamstead by Mr H. Insley and at 
Sandwell by Mr H. W. Hughes, F.G.S. 

Horizon and Localities. — Blue Measures, six feet above Brooch Coal : Hamstead 
Colliery, Great Barr, near Birmingham ; Jubilee Pit, Sandwell Park, West Bromwich. 

Sphenopteris Souichi Zeiller. 

1886. Sphenopteris Souichi, Zeiller, Flore foss. bassin houil. d. Valen., p. 110, pi. vii. figs. 2, 2a, 2b, 

pi. ix. figs. 3, 3a, 3b. 

Remarks.— I am indebted to Dr Wheelton Hind for an example of this species. 
The fossil is preserved in an ironstone nodule, and, as it is somewhat fragmentary, the 
specimen was forwarded to Dr Zeiller, who has kindly compared it with the type, 
with which he says it agrees perfectly. 

Sphenopteris Souichi has not previously been recorded for Britain. 

Horizon and Locality. — Ten foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

* Uber einige Carbonfarne aus der Familie der Sphenopteriden : Inaugural Dissertation, p. 37, 1908. 

t Gothan, "Die Oberschlesische Steinkohlenfiora. I. Theil: Fame und famahnliche Gewachse," p. 139, 
Abhandl. d. konig. preuss. geol. Landesanstalt, neue Folge, Heft 75, 1913. 

\ Fame d. Carbon-Flora d. Schatzlarer Schichten, p. 58, pi. xxxix. fig. 6, text-fig. 12a, b, 1885. 

5; Stur, Carbon-Flora d. Schatzlarer Schichten: Die Fame, p. 58, pi. xxxix. figs. 7, la ; pi. xl. figs. 2-6 (not fig. 1, 
1885). 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 89 



Sphenopteris Schatzlarensis Stur, sp. 

1885. Hapalopteris Schatzlarensis, Stur (pars), Carbon-Flora d. Scliatz. Schichten : Die Fame, p. 58, 

pi. xl. figs. 2-6, pi. xxxix. figs. 7 and la, and text-fig. 11, p. 59 (non pi. xl. fig. 1). 
1899. Sphenopteris (Renaultia) Schatzlarensis, Zeiller, Flore foss. bassin houil. d. Heraclee, p. 15, 

pi. i. fig. 11. 

1910. ,, ,, ,, Deltenre, in Renier, Documents paleont. terr. houil., 

pi. lxiii. 
1913. Sphenopteris (? Renaultia, ? Boweria) Schatzlarensis, Gothan, Oberschlesische Steinkohlenflora, 

i. Theil, p. 139, pi. xxviii. fig. 5, pi. xxix. figs. 1 and 4. 
1908. Ovopteridium Schatzlarense, Behrend, Uber einige Garbonfarne der Familie d. Sphenopteriden : 

Inaugural Dissertation, p. 37. (Also in Jahrb. d. k. preuss. geol. Landesanstalt, vol. xxix., 

L, Heft iii. p. 679.) 

1911. Boweria Schatzlarensis, Kidston (pars), Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. 

p. 34. 

1885. Hapalopteris Aschenborni, Stur, I.e., p. 63, pi. xxxix. fig. 6 (text-figs. 12a and 12/; inaccurate). 
1899. Ovopteris Aschenborni, Potonie, Lehrb. d. Pflanzenpal., p. 143, fig. 136 (copied from Stur). 

Remarks. — For notes on this species, see Boweria Schatzlarensis, p. 90. 
Horizons and Localities. — 

Roof of Neu) Mine Coal: Clattershall Colliery, Brettell Lane. 

Blue Measures, six feet above Fireclay Coal : Doulton's Clay Pit, Netherton, 
near Dudley. 

Renaultia Zeiller. 
Renaultia gracilis Brongniart, sp. 

1829. Sphenopteris gracilis, Brongt., Hist. d. veget. foss., vol. i. p. 197, pi. liv. fig. 2. 

1870. „ ,, Lesqx., Geol. Surv. of lllin., vol. iv. p. 408 (non pi. xv. figs. 3-6). 

1883. „ „ Renault, Cours d. bot. foss., vol. iii. p. 189, pi. xxxiii. figs. 1-4. 

1886. Sphenopteris (Renaultia) gracilis, Zeiller, Flore foss. bassin houil. d. Valen., p. 94, pi. iv. figs. 2, 3. 

1912. ,, ,, ,, Gothan, Verhandl. d. naturhist. Vereins d. preuss. Rheinl. u. 
Westph., 69. Jahrgang, p. 248, pi. iii. figs. 3-5. 

1829. Sphenopteris fragilis, Brongt., Hist. d. veget. foss., pi. liv. fig. 2. 

1882. Sphenopteris microcarpa, Kidston (non Lesqx.) (pars), Proceed. Roy. Phys. Soc, vol. vii. p, 129, 

pi. i. figs. 9-14 (non figs. 7, 8). 
1888. Renaultia microcarpa, Zeiller, Flore foss. bassin houil. d. Valen., p. 29, fig. 15. 

Horizons and Localities. — 

Roof of Thick Coal: Bradley Colliery, Bilston. 

Roof of New Mine Clay : Clattershall Colliery, Brettell Lane. 

Roof of New Mine Coal: Mount Pleasant, Brierley Hill; Clattershall Colliery, 

Brettell Lane. 
Roof of Fireclay Coal : Doulton's Clay Pit, Netherton, near Dudley. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 

Boweria Kidston. 

1911. Boweria, Kidaton, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 31. 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 12 



90 DR ROBERT KIDSTON ON THE 

Boweria Schatzlarensis Kidston. 

1885. Hapalopteris Schatzlarensis, Stur (pars), Carbon-Flora d. Schatz. Schichten: Die Fame, p. 58, 

pi. xl. fig. 1 (non figs. 2-6). 
1911. Boweria Schatzlarensis, Kidston, Mem. Musee roy. d'hist. not. de Belgique, vol. iv. p. 34 

(refs. in part), text-figs. 5, 6, p. 32. 
1890. Renaultia Schatzlarensis, Kidston, Trans. York. Nat. Union, part xiv. p. 32. 
1890. Cf. Renaultia (Sphenopteris) microcarpa, Potonie\ " Ueber einige Carbonfarne," part i., Jahrb. 

d. It. preitss. geol. Landesanstalt fitr 1889, p. 25, pi. v. figs. 3, 3a. 
1869. Sphenopteris Bronni, Roehl (non Gutbier), Foss. Flora Steink.-Form. Westphalens, p. 57, pi. xvi. 

figs. 5, 5a. 

Remarks. — Under the name of Hapalopteris Schatzlarensis Stur included two 
species. This has been pointed out by Behrend * and Gothan. That given at fig. 1, 
pi. xl. by Stur is specifically distinct from the other specimens included by him 
under the same name. This fig. 1 forms the type of the genus Boweria, whose 
fructification consists of small annulate, circular or oval sporangia, free, one placed on each 
veinlet at the margin of the pinnule, and measuring about 0"40 mm. in their greatest 
diameter. The annulus forms a band of two rows of prominent cells which passes 
across the apex of the sporangium and extends a very short distance down the sides. 
The limb of the fertile pinnules is slightly reduced, the segments being obtuse or 
truncate and not pointed as in the sterile condition. The other of the two species 
included by Stur under the same name is placed in the genus Sphenopteris under the 
specific name given it by Stur — Sphenopteris Schatzlarensis. 

Boweria Schatzlarensis Kidston and Sphenopteris Schatzlarensis Stur seem to 
have a similar distribution. Both occur in the South Staffordshire Coal Field, but are 
not common. They also both occur in the Yorkshire Coal Field, where, however, 
Boweria Schatzlarensis appears to be the more common of the two. 

I also possess a small fertile specimen (No. 3541) of Boweria Schatzlarensis, 
received a few years ago from M. L. Deltenre, Mariemont, from Ste. Henriette Pit, 
on which the annulus of the sporangia is well shown. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

Crossotheca Zeiller. 
Crossotheca Honinghausi Brongt., sp. 

1829. Sphenopteris Honinghausi, Brongt., Hist. d. veget. foss., vol. i. p. 199, pi. Hi. 
1865. ,, ,, Andrae, Vorwelt. Pflanzen, p. 13, pis. iv., v. 

1905. Crossotheca Honinghausi, Kidston, Proc. Roy. Soc, ser. B, vol. lxxvi. p. 358, pi. vi. figs. 1-5. 

1906. „ „ Kidston, Phil. Trans., ser. B, vol. cxcviii. p. 413, pi. xxv. figs. 1-16, 

pi. xxvi. figs. 17-32, text-figs. 1-7. 

* Behrend, Uber einige, Carbonfarne aus der Familie der Rphenopteriden : Inaugural Dissertation, 1908, p. 37. 
Also in Jahrb. d. k. preuss. geol. Landesanstalt, vol. xxix., I., Heft iii. p. 679 ; Gothan, Oberscldesische Steinkohlenflora 
i. Theil, p. 139, 1913. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 91 

Note. — Frequent. 
Horizons and Localities. — 

Roof of Brooch Coal: Pensnett. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley ; 
Ettingshall. 

Roof of Thick Coal : Bradley Colliery, Bilston. 

Roof of Fireclay Coal : Doulton's Clay Pit, Netherton. 

Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 

Immediately below Bottom Coal : Kuiton, near Sedgley. 

Crossotheca Hughesiana Kidston. 

1906. Crossotheca Hughesiana, Kidston, Phil. Trans., ser. B, vol. cxcviii. p. 424, pis. xxvii.-xxviii. 

Horizon and Locality. — Ten-foot Ironstone Measures ( = Roof of Thick Coal): 
Clayscroft Openwork, Coseley, near Dudley. 

Crossotheca Crepini Zeiller. 

1883. Crossotheca Crepini, Zeiller, Ann. d. Soc. not., 6 e s^r., Bot., vol. xvi. p. 181, pi. ix. figs. 1-9. 
1886. Sphenopteris (Crossotheca) Crepini, Zeiller, Flore foss. bassin houil. d. Valen., p. 112, pi. xiii. 

figs. 1-3; text-fig. 21, p. 33. 
1910. Sphenopteris (Crossotheca) Crepini, Schmitz, in Renier, Paleontologie, pi. Ixviii. 
1883. Sorotheca Crepini, Stur, Zur Morph. u. Syst. d. Culm- u. Carbonfarne, p. 175 (text-fig. 39 

not good). 
1885. Sorotheca Crepini, Stur, Carbon-Flora d. Schatz. Schichten : Die Fame, p. 275, pi. xxxiii. figs. 1 

and 2, pi. xxxv. figs. 3-4 (text-fig. 43, p. 273, not good). 

Remarks. — This species, which has not previously been recorded from Britain, has 
been collected by Mr H. W. Hughes at Grets Green and by Mr H. Insley at 
Hamstead. 

Horizon and Localities. — 

Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr, 

near Birmingham. 
Roof of Brooch Coal : Grets Green, near West Bromwich. 

Sphyropteris Stur. 
Sphyropteris obliqua Marrat, sp. 

1872. Sphenopteris obliqua, Marrat, in Higgins, Proc. Liverpool Geol. Soc, Session 13, 1871-1872, 

p. 99, pi. ix. figs. 3, 3a. 
1889. Sphyropteris obliqua, Kidston, Trans. Roy. Soc. Edin., vol. xxxv. p. 402, pi. i. figs. 3, 3a-3a\ 
1883. Sphyropteris Crepini, Stur, Die Fame: Carbon-Flora d. Schatzlarer Schicht., p. 18, pi. xxxix. 

figs. 1, la; text-fig. 6a, p. 16. 



92 DR ROBERT KIDSTON ON THE 

Horizon and Localities. — 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Roof of New Mine Coal: Mount Pleasant, Brierley Hill. 
Roof of Fireclay Coal: Doulton's Clay Pit, Netherton. 

Zeilleria Kidston. 
Zeilleria Avoldensis Stur., sp. 

PL VII. figs. 5, 5a, 5b, G, 6a. 

1878. Fhtinophyllum Avoldense, Stur, Verh. d. k. k. geol. Reichsanstalt, p. 213. 

1883. Calymmotheca Avoldensis, Stur, "Morph. u. Syst. d. Culm- u. Carbonfarne," Sitzb. d. k. Akad. d. 

Wissensch., vol. lxxxviii. p. 171, fig. 37. 
1885. ,, ,, Stur, Fame d. Carbon-Flora d. Scliatzlarer Schichten, p. 251, 

pi. xxxvii. fig. 1, text-fig. 41, p. 238. 
1899. ,, „ Potonie\, Lehrb. d. Pflanzenpal., p. 103, fig. 90 i. 

1884. Zeilleria Avoldensis, Kidston, Quart. Journ. Geol. Soc, vol. xl. p. 591. 

1887. ,, ,, Kidston, Trans. Roy. Soc. Edin., vol. xxxiii. p. 148, pi. viii. figs. 8-10. 

Description. — Frond very large quadripinnate. Primary pinnae broadly 
lanceolate ; secondary pinnae alternate, linear-lanceolate ; tertiary pinnae alternate, 
linear-lanceolate, or oblong lanceolate. Pinnules alternate, attached by their whole 
base to the rachis and united to each other for one-third to two-thirds of their length ; 
free portion of limb ovate triangular. Nervation clearly defined, and consisting of a 
central vein which generally gives off a lateral veinlet on each side, which extends to 
the margin of the pinnule. 

Fructification apparently confined to the tertiary pinnse on the lower secondary 
pinnse, where the fertile pinnules, similar in form to the sterile ones, bear one to three 
pedicellate cupule-like structures placed at the extremities of the excurrent veins. 
When immature the fructification appears as a closed oval structure, but at maturity 
it splits into four, rarely into five, spreading segments, which on the specimens with 
larger cupules show in the centre a small mammillate point to which a small organ, 
presumably a seed, has been attached. What are probably the microsporangia are 
very similar in form to the seed-bearing cupules when unopened, but are of smaller size. 

Remarks. — Two specimens of this fern are given on PI. VII. figs. 5, 6. That given 
natural size at fig. 6 represents fairly well the general character of a fertile specimen, 
though the structure of the fructification is not well exhibited. A small portion of 
the same specimen is shown enlarged 2 times at fig. 6a. Most of the sporangia seem 
to have adhered to the counterpart of the specimen, which unfortunately has not been 
preserved. They are much better seen on another small example, also from the South 
Staffordshire Coal Field, which has already been figured in vol. xxxiii., pi. viii. figs. 
8-10, of the Transactions of this Society. Here the small cupule-like structures are 
seen to be oval when immature, but split at maturity into four valves about 1 mm. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 93 

long, text-fig. 2. Another specimen, preserved in an ironstone nodule, is shown 
natural size at fig. 5, and, although the specimen is somewhat broken up, the structural 
details of the cupules are very beautifully exhibited, and can be seen with a lens 
on the part of the specimen given enlarged about 2\ times at fig. 5a. A further 
enlargement of the cupule lettered a on fig. 5a is seen at fig. 5b, where it is enlarged 
about 6 times. The spread-out cupules on this example measure about 3 mm. across, 
and are therefore larger than on the specimens to which reference has been made 
above. They further show most distinctly a central prominent mammillate point, to 
which I have no doubt a small seed was originally attached, though now fallen off. 
To show that this is not an improbable explanation, it may be mentioned that small 
Carpolithes-like seeds, only 1*5 mm. long and about 1 mm. wide, have been found in 
the Lanarkian Series of Midlothian by Mr D. Tait, which when treated by the 
maceration process showed the nucellus, pollen-chamber, and microspores, hence the 
cupule was large enough to enclose such a seed. 





b 
Text-jig. 2. —Zeillcria Avoldensis Stur, sp. a shows sporangia open, and b immature, x 3J times. 

Each of the segments of these larger cupules has a much more dense central 
portion, as indicated by the amount of carbonaceous matter still preserved, and a 
narrow, more delicate, margin. This is seen at fig. 5b, which shows a cupule enlarged 
about 6 times. Possibly the thin marginal bands may represent overlapping portions 
of the segments of the cupule. 

To return to the specimen described on PI. VIII. figs. 8-10 of vol. xxxiii. of the 
Transactions of this Society, the cupule-like structures, as already mentioned, are 
smaller than those given at fig. 5, and which are supposed to have enclosed a seed. 
Text-fig. 2 shows two pinnules from the earlier specimen enlarged about 3|- times ; 
that at a shows the attached bodies split into four valves, while that at b, 
from the same specimen, shows a single oval unopened body attached to its 
short stalk. Fructifications similar in appearance to this latter were described by 
Dr Arber as the seeds of a Pteridosperm, under the name of Carpolithus Nathorsti* 
Others of his figures f seem to show the structure divided into valves similar to the 
figure given here. Subsequently Nathorst showed that the bodies which had been 
regarded as Carpolithus by Dr Arber were really sporangia containing microspores, J 

* Ann. of Bot., vol. xxii. p. 57, pi. vi. fig. 4. 
t Loc. cit., figs. 3, 5, and 6. 

\ Nathorst, Palaeobot. Mitteilungen, No. 4 ; Kungl. Svenska Vetenskaps-Akad. Handl., Band xliii. No. 6, p. 10, 
pi. ii. tigs. 19-21. 



94 DR ROBERT KIDSTON ON THE 

and that the specimen which had been described as Carpolithus Nathorsti was in 
all probability a fertile example of Zeilleria Schaumburg-Lippeana Stur, sp. # These 
discoveries therefore make it extremely probable that the specimen of Zeilleria 
Avoldensis given at text-fig. 2 represents the microsporangia-bearing plant, and that 
shown at PI. VII. fig. 5 the seed plant. The larger size of the split cupule of fig. 5 
cannot be regarded as merely a stronger development, and the central mammillate 
protuberance can scarcely be explained otherwise than the point of attachment of a 
small seed. 

The form of the pinnules bearing these larger and smaller fructifications is similar. 
The specimens of Zeilleria Avoldensis, sp., figured here were collected by 
Mr H. W. Hughes, who kindly placed them in my hands for description. 
Horizon and Localities. — 

Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr, 

near Birmingham. Collected by Mr H. Insley. 
Roof of Brooch Coal : Shut End ; Tividale. 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 



Zeilleria delicatula Sternberg, sp. 

1823. Sphenopteris delicatula, Sternb., Essai flore monde prim., vol. i. fasc. ii. p. 34, pi. xxvi. fig. 5 ; 

fase. iv. p. 16. 
1829. „ „ Brongt., Hist. d. veget. foss., p. 185 (?pl. lviii. fig. 4). 

] 848. ,, „ Sauveur, Veget. foss. terr. houil. de la Belgique, pi. xxiii. fig. 5. 

1899. ,, ,, Hofmann et Ryba, Leitpjianzen, p. 42 (? pi. iii. figs. 22, 22a). 

1884. Zeilleria delicatula, Kidston, Quart. Journ. Geol. Soc, vol. xl. p. 592, pi. xxv. figs. 1-12. 
1888. ,, ,, Zeiller, Flore foss. bassin houil. de Valen., p. 57, fig. 37 a, b. 

1891. „ ,, Kidston, Trans. Geol. Soc. Glasgow, vol. ix. p. 33, pi. iii. fig. 34 a, b. 

1899. ,, ,, Potonie, Lehrb. d. PJlanzenpal., p. 103, fig. 90 in. 

1903. ,, „ Arber, Quart. Journ. Geol. Soc, vol. lix. p. 13, pi. ii. figs. 1, 2. 

Remarks. — Both sterile and fertile states of the plant have been collected — those 
at Sandwell by Mr H. W. Hughes, and those at Hamstead by Mr Henry Insley. 

Horizon and Localities. — Blue Measures, above Brooch Coal : Jubilee Pit, Sandwell 
Park, West Bromwich ; Hamstead Colliery, Great Barr, near Birmingham. 



Zeilleria sp. 

Remarks. — Mr H. W. Hughes has collected some fragments of an undescribed 
species of Zeilleria at Sandwell ; but as I have better specimens of this plant from the 
Yorkshire and Burnley Coal Fields, I shall defer its description until another time. 

Horizon and Locality. — Blue Measures, above Brooch Coal : Jubilee Pit, Sandwell 
Park, West Bromwich. 

* See Nathorst, I.e., p. 10 ; and Zeiller, Comptes rendus, vol. cxliv. p. 1139, 1907. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 95 

Eremopteris Schimper. 
Eremopteris artemisiaefolia Sternb., sp. 

1826. Sphenopteris artemisieefolia, Sternb., Essai flore monde prim., vol. i. fasc. iv. p. xv. pi. liv. fig. 1. 
1829. „ „ Brongt., Hist. d. veget. foss., p. 176, pi. xlvi., pi. xlvii. figs. 1, 2. 

1869. Eremopteris artemisixfolia, Schimper, Traite d. paleont. veget., vol. i. p. 416, pi. xxx. fig. 5. 
1877. Sphenopteris (Eremopteris) artemisixfolia, (?) var. Lebour, Illustr. of Fossil Plants, pi. xxxiii. 
1826. Sphenopteris stricta, Sternb., Essai flore monde prim., vol. i. fasc. iv. p. xv, pi. lvi. fig. 3; vol. ii. 

p. 57. 
1829. ,, ,, Brongt., Hist. d. veget. foss., p. 208, pi. xlviii. fig. 2. 

1832. Sphenopteris crithmifolia, L. & H., Fossil Flora, vol. i., pi. xlvi. 
1877. Sphenopteris sp., Lebour, Illustr. of Fossil Plants, pis. xxxiv., xxxv., xxxvi. 
Asplenioides obtusum, Konig, Icones foss. sectiles, pi. xvi. fig. 199. 

Horizon and Locality. — Blue Measures, six feet above Fireclay Coal: Doulton's 
Clay Pit, Netherton. 

Adiantites Goppert. 
Adiantites sessilis Roehl, pro. var. 

1869. Cyclopteris oblongifolia, Gopp., var. sessilis, Roehl, Foss. Flora Steink.-Form. Westphal., p. 45, 

pi. xvi. figs. 1 and la. 
1899. Adiantites sessilis, Potonie, Lehrb. d. Pflanzenpal., p. 129, fig. 116. 

Remarks. — Of this species I have only seen two British specimens — that noted here, 
and one from Adderley Green, Longton, North Staffordshire, from " below the Bowling- 
alley Rock," which was received from the late Mr John Ward, F.G.S. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. Collected by Mr H. W. Hughes. 

Archseopteris Dawson. 

Archeeopteris Reussi Ettingshausen, sp. 
PL V. figs. 7 and 7a. 

1852. Asplenites Reussii, Ett., "Steinkf. v. Stradonitz in Bohmen," Abhandl. d. h. geol. Eeichsanst., 

I., Band iii. Abth. No. 4, p. 16, pi. i. figs. 8, 9. 
1869. Palxopteris Reussii, Schimper, Traite d. paleont. veget., vol. i. p. 478. 
1894. Archseopteris Reussii, Kidston, Proc. Roy. Phys. Soc. Edin., vol. xii. p. 242. 

Description. — Frond bipinnate, main rachis rigid, straight, smooth ; pinnse alternate, 
linear-lanceolate, rachis straight, smooth ; pinnules alternate, obovate, directed forwards 
and contracted into a short decurrent stalk, upper margin minutely dentate ; nervation 
flabelliform, arising from the repeated dichotomy of a single vein. 

Remarks. — The pinnules are so directed forwards that their margin next the rachis 
lies almost parallel with it. They are gradually narrowed into a short stalk, into which 



96 DR ROBERT KIDSTON ON THE 

a single vein enters, and, dichotomising four or five times in its course, provides from 
sixteen to over twenty ultimate veinlets, one of which seems to enter each little tooth 
on the upper margin of the pinnule, but the little teeth are seldom well seen on our 
specimen, which has suffered somewhat from decay before fossilisation took place. 

ArchsBopteris Reussi is described as bipinnate, but possibly the plant attained a 
greater size and only fragments of lateral pinnse may be known. I have compared the 
British example with a small specimen of Archseopteris Reussi from Stradonitz in the 
British Museum, and find it agrees with it perfectly. 

Hitherto Archseopteris Reussi Ett., sp., appears only to have been found at 
Stradonitz, Bohemia. 

In the absence of fructification some slight doubt may exist as to whether 
Archseopteris Reussi should be retained in Archseopteris, though its foliage pinnules 
agree with that genus in every respect. 

As the only other course would be the creation of a new genus for its reception, and 
that only distinguished by negative characters, I retain it provisionally in Archseopteris 
till some knowledge of its fructification is obtained. In any case it is a remarkable 
circumstance that plants possessing the foliage of the genus Archseopteris, of the 
Archseopteris hibernica Forbes, sp., type, should disappear from all the intervening 
rocks that lie between the Upper Old Red Sandstone and the Westphalian Series of the 
Upper Carboniferous, and should after this great length of time again occur without 
any apparent generic alteration. 

Archseopteris Reussi seems to have been a smaller species than any of the Upper 
Old Red Sandstone members of the genus. 

The specimen from Tividale is shown natural size at fig. 7, and some pinnules 
enlarged 1\ times are given at fig. 7a. 

The fossil is contained in the " Johnson Collection " of the Department of Geology 
and Palaeontology of the British Museum, No. V., 1366, and my thanks are due to 
Dr A. Smith Woodward, F.R.S., for kind permission to figure an<7 describe it. 

Horizon and Localities. — 

Blue Measures above Brooch Coal: Jubilee Pit, Sandwell Park, West 
Bromwich. Collected by Mr. H. W. Hughes. Hamstead Colliery, Great 
Barr, near Birmingham. Collected by Mr Henry Insley. 
Westphalian Series : Tividale, near Dudley, Staffordshire. 

Telangium Benson. 
Telangium asteroides Lesqx., sp. 

1870. Staphylo[deris asteroides, Lesqx., Oeol. Survey, of Illin., vol. iv. p. 406, pi. xiv. figs. 6, 7 (non 

figs. 8-10). 
1879. Sorocladus asteroides, Lesqx., Coal Flora, p. 328, pi. xlviii. figs. 9, 9a, and 9b. 
1883. Calymmatotheca asteroides, Zeiller, Ann. d. sc. nat., 6 e s6r., Bot., vol. xvi. pp. 182 and 207, pi. ix. 

figs. 10, 11. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 97 

1886. Calymmatotheca asteroides, Zeiller, Flore foss. bassin houil. d. Valen., p. 141, pi. xii. figs, 2, 2a, 

and 2b. 
1891. „ „ Kidston, Trans. Geul. Soc. Glasgow, vol. ix. p. 24, pi. iii. fig. 37. 

Remarks. — A number of years ago I saw a small specimen of this species in the 
collection of the late Mr Henry Johnson, Dudley. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

Coseleya Kidston, n.g. 

Description. — Sporangia pyriform, exannulate, sessile, attached spirally or in close 
verticils round the rachis, and forming dense masses. 



Coseleya glomerata Kidston, n. sp. 
PI. V. figs. 4, 4a, 5, and 6 ; PL X. fig. 4. 

Description. — Rachis stout, slightly flexuous, with closely placed, irregular, small 
transverse scars ; pinnae alternate, short, and entirely obscured by closely placed 
sporangia arranged spirally or in whorls around the rachis. Sporangia pyriform, 
exannulate, pedicellate, and free. Apparently dehiscing by a longitudinal cleft on 
their ventral surface. 

Remarks. — At fig. 4, PI. V., the largest specimen of this fructification which has 
been discovered is shown natural size. The sporangia themselves have decayed, and 
the groups only leave their impression in the matrix, accentuated by a dark brown 
stain. The same specimen is shown enlarged 2 times at fig. 4a. At the part- 
marked a on this figure, one of the pinnae has been broken over and shows the groups 
of sporangia in section, where at one point the form of the sporangia can be clearly 
made out. Two of these are enlarged 9 times at fig. 46. They are pyriform, and 
their base is contracted into a short pedicel. 

Occasionally one sees a dark line running down the ventral face of the sporangia, 
which probably indicates the position of a longitudinal cleft by which dehiscence 
took place. 

The rachis is stout, but this was necessary for the support of the dense masses of 
sporangia which the short pinnae bore. The main rachis of this specimen must have 
been densely clothed with scales, or hairs, from the numerous scars they have left to 
indicate their former presence. 

At figs. 5 and 6 two other specimens of the same species are given. At a on fig. 6 
the arrangement of the sporangia around the axis is also well seen. It is difficult to 
determine whether the sporangia are in whorls or arranged spirally on the rachis, but 
I rather think they are spirally placed. At the upper part of fig. 5 the form of the 
sporangia is shown ; but the sporangia occur in such dense masses that one sporangium 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 13 



98 DR ROBERT KIDSTON ON THE 

generally tends to obscure the form of the other, and the general appearance of the 
pinrme is as if the stem bore small bramble-like fruits, the " pips" being the impressions 
of the apices of the sporangia in the matrix, as seen at fig. 4, PI. X., which is enlarged 
2 times. 

Possibly in Coseleya glomerata we see the microsporangia of a pteridosperm, but 
it is impossible from the meagre data at present at our disposal to form any definite 
opinion of the affinities of Coseleya. 

The genus name is derived from Coseley, near Dudley, where in the past so many 
fine specimens have been obtained. 

All the specimens of Coseleya glomerata which I have seen were collected by 
Mr H. W. Hughes, but it is evidently a rare fossil. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Pecopteris Brongniart. 
Pecopteris Miltoni Artis, sp. 

1825. Filicites Miltoni, Artis, Antedil. Phyt., pi. xiv. 

1834. Pecopteris Miltoni, Brongt. (pars), Hist. d. veget. foss., p. 333, pi. cxiv. fig. 8 (non figs. 1-7). 
1849. ,, ,, Andrae, in Germar, Vers. d. Steinh. v. Wettin u. Lbbejun, p. 63, pi. xxvii. 

(Excl. syn., Pecopteris polymorpha. Refs. in part.) 
1911. Pecopteris (Asterotheca) Miltoni, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. 
p. 50. 

1885. Hawlea Miltoni, Stur, Carbon-Flora d. Schatzlarer Schicht. ; Die Fame, p. 108, pi. lix. figs. 1-4; 

pi. lx. figs. 1, 2 (excl. figs. 3, 4) ; text-fig. 17, p. 106 (syn. in part). 

1835. Pecopteris abbreviata, Brongt., Hist. d. veget. foss., p. 337, pi. cxv. figs. 1-4. 

1836. „ „ L. & H., Fossil Flora, vol. iii., pi. clxxxiv. 

1886. Pecopteris (Asterotheca) abbreviata, Zeiller, Flore foss. bassin houil. d. Valen., p. 186, pi. xxiv. 

figs. 1-4. 

Note. — Frequent. 
Horizons and Localities. — 

(?) Blue Measures above Brooch Coal : Jubilee Pit, Sandwell Park, West 
Bromwich. 

Blue Measures, six feet above Brooch Coal: Hamstead Colliery, Great Barr, 
near Birmingham. 

Brooch Coal: Oldbury. 

Ten-foot Ironstone Measures: Clayscroft Openwork, Coseley, near Dudley. 

Roof of Thick Coal: Bradley Colliery, Bilston. 

Roof of Neiv Mine Coal: Coseley; Clattershall Colliery, Brettell Lane. 

Six feet above Fireclay Coal: Doulton's Clay Pit, Netherton. 

Between Fireclay and Bottom Coals : Doulton's Clay Pit, Netherton. 

Immediately beloiv Bottom Coal: Ruiton, near Sedgley. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 99 

Pecopteris Volkmanni Sauveur. 

1848. Pecopteris Volkmanni, Sauveur, Veget. foss. terr. houil. d. Belgique, pi. xlv. figs. 1-4. 
1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 204, pi. xxviii. figs. 1-3. 

1882. Cyatheites arborescens, Achepohl {non Schlotheim), Niederrh. Westfdl. Steink., p. 94, pi. xxxii. fig. 1. 

1885. Senftenbergia Boulay, Stur, Carbon-Flora d. Schatzlarer Schichten : Die Fame, p. 85, pi. 1. fig. 1. 

Note.— Not common. 
Horizons and Localities. — 

Blue Measures above Brooch Coal : Jubilee Pit, Sandwell Park, West 
Bromwich. 

Roof of New Mine Coal: Mount Pleasant, Brierley Hill. 

Blue Measures, six feet above Fireclay Coal: Doulton's Clay Pit, Netherton. 

Pecopteris ? hepaticseformis Kidston, n. sp. 
PI. VII. fig. 7. 

Remarks. — -A figure of a fragment of the smallest pinnuled fossil fern I have yet 
seen is given natural size on PI. VII. fig. 7. The portion preserved shows part of a 
pinna about 11 mm. long and about 1"5 mm. wide at its base, from where it decreases 
very gradually upwards. The specimen is converted into a very black carbonaceous 
substance, and indicates that the pinnules must have been of considerable thickness. 
The nervation is not distinctly seen. Two of the pinnules, magnified 10 times, are 
shown at fig. 7a, but some of them appeared to be slightly more lobed than those 
enlarged. I have met with no fossil at all similar to this curious H epaticse-Wke little 
species, which, however, in the absence of more complete specimens, I have placed 
provisionally amongst the Pecopterids. 

Horizon and Locality. — Ten-foot Ironstone Measures : Coseley (?). In the 
collection of Sir Charles Holcroft. 

Dactylotheca Zeiller. 
Dactylotheca plumosa Artis, sp. 

1825. Filicites plumosus, Artis, Antedil. Phyt., p. 17, pi. xvii. 

1836. Pecopteris plumosa, Brongt., Hist. d. veget. foss., p. 348, pis. cxxi., cxxii. 

1885. Senftenbergia plumosa, Stur, Carbon-Flora d. Schatzlarer Schichten: Die Fame, p. 92, pi. li. figs. 1-3. 

1896. Dactylotheca plumosa, Kidston, Trans. Roy. Soc. Edin., vol. xxxviii. p. 205, pis. i.-iii. 

1834. Sphenopteris crenata, L. & H, Fossil Flora, p. 59, pis. c, ci. 

1885. Senftenbergia crenata, Stur, Carbon-Flora d. Schatzlarer Schichten: Die Fame, p. 72, pi. xlv. 

figs. 1-3, pi. xlvi. figs. 1-3. 
1834. Schizopteris adnascens, L. & H., Fossil Flora, vol. i. p. 58, pis c. and ci. 
1832. Sphenopteris caudata, L. & H., Fossil Flora, vol. i. p. 137, pi. xlviii.; vol. ii. p. 157, pi. cxxxviii. 

1834. Pecopteris dentata, Brongt., Hist. d. veget. foss., p. 346, pis. cxxiii., cxxiv. 

1835. „ „ L. & H., Fossil Flora vol. ii. p. 201, pi. cliv. 

1886. Pecopteris {Dactylotheca) dentata, Zeiller, Flore foss. bassin houil. d. Valen., p. 196, pi. xxvi. 

figs. 1, 2 ; pi. xxvii. figs. 1-4 ; pi. xxviii. figs. 4, 5. 
1899. „ ,, „ White, Foss. Flora Low. Coal Meas. of Missouri, p. 75, pi. xxiv. 

figs, 1, 2 ; pi. xxv.; pi. xxvi. figs. 2-4; pi. xxvii. 



100 DR ROBERT KIDSTON ON THE 

1836. Aspidites silesiacus, Gopp., Syst. fil.foss., p. 364, pi. xxvii. (pi. xxxix. fig. 1?). 

1836. Pecopteris delicatula, Brongt., Hist. d. veget. foss., p. 349, pi. cxvi. fig. 6. 

1854. Pecopteris Glorkeriana, Ett. (Gopp. 1), St ink/, v. Radnitz, p. 44, pi. xvii. fig. 1. 

1854. Pecopteris angustifida, Ett., Stinkf. v. Radnitz, p. 45, pi. xvi. fig. 1. 

Note. — This species is widely distributed in the South Staffordshire Coal Field, 
where it is common. 

Horizons and Localities. — 

Blue Measures above Brooch Coal: Jubilee Pit, Sandwell Park, West 

Bromwich. 
Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr, 

near Birmingham. 
Above Brooch Coal: Grets Green, near West Bromwich. 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Roof of Thick Coal: Bradley Colliery, Bilston. 
Roof of New Mine Coal: Coseley, near Dudley ; Clattershall Colliery, Brettell 

Lane. 
Blue Measures, six feet above Fireclay Coal: Doulton's Clay Pit, Netherton. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 
Five yards above White Ironstone : No. 19 Pit, Saltwells, Cradley. 
Roof of Bottom Coal: Bradley. 

forma delicatula Brongt., pro. sp. 

Above Brooch Coal: Oldbury. 

Roof of Fireclay Coal: Russell's Hall, Dudley. 

Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 

Mariopteris Zeiller. 
Mariopteris muricata Schl., sp. 

1804. Schlotheim, Flora d. Vorwelt, pp. 54, 55, pi. xii. figs. 21 and 23. 

1820. Filicites muricatus, Schloth., Petrefactenkunde, p. 409. 

1832. Pecopteris muricata, Brongt., Hist. d. veget. foss., p. 352, pi. xcv. figs. 3, 4, pi. xcvii. 

1885. Diplothmema muricatum, Stur, Carbon-Flora d. Schatz. Schichien : Die Fame, p. 395, pi. xxi. 

figs. 1-5, pi. xxii. figs. 1-5, pi. xxiii. figs. 1-6. 
1879. Mariopteris muricata, Zeiller, Bull. Soc. geol. de France, 3 e ser., vol. vii. p. 92. 
1886 - .» >» Zeiller, Flore foss. bassin houil. d. Valen., p. 173, pi. xx. figs. 2, 3; pi. xxi. 

fig. 1 ; pi. xxii. fig. 2. 
1^11- „ „ Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 53. 

1886 - » ., forma nervosa, Zeiller, ibid., pi. xx. fig. 1, pi. xxii. fig. 1, pi. xxxiii. fig. 1. 

1886 - >. „ var. hirta, Zeiller, ibid., p. 182, pi. xx. fig. 4. 

1879. Psendo-pecopteris muricata, Lesqx., Coal Flora, vol. i. p. 203, pi. xxxvii. fig. 2. 

1832. Pecopteris nervosa, Brongt., Hist. d. veget. foss. p. 297, pi. xciv., pi. xcv. figs. 1, 2. 

1833. ,, „ L. & H., Fossil Flora, vol. ii. pi. xciv. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 101 

1885. Diplothmema nervosum, Stur, Carbon-Flora d. Schatz. Schichten: Die Fame, p. 384, pi. xxiv. 

fig. 1 ; pi. xxv. b., fig. 2. 
1834. Pecopteris laciniata, L. & H., Fossil Flora, vol. ii. pi. cxxii. 

1885. Diplothmema hirtum, Stur, ibid., p. 372, pi. xxxiv. fig. 1. 

Note. — Very common, and widely distributed in the coal field. 
Horizons and Localities. — 

Roof of Brooch Coal; Pensnett; Himley; Shut End; Kingswinford ; Foxyards, 

Dudley. 
Blue Measures above Brooch Coal : Jubilee Pit, Sand well Park, West Bromwich ; 

Hamstead Colliery, Great Barr, near Birmingham. 
Roof of Thick Coal: Bradley Pit, Bilston. 

Ten-foot Ironstone Measures: Clayscroft Openwork, Coseley, near Dudley. 
Roof of New Mine Coal : Mount Pleasant, Brierley Hill ; Doulton's Clay Pit, 

Netherton. 
Roof of Bottom Coal: No. 120 Pit, Coneygre, Tipton. 

forma nervosa Brongt, pro. sp. 

Ten foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Shales, thirty yards above Thick Coal : Hamstead Colliery, Great Barr, near 

Birmingham. 
Shale, over Thick Coal : Near Bilston. 
Roof of New Mine: Coseley, near Dudley. 
Roof of Fireclay Coal: Doulton's Clay Pit, Netherton. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 
Immediately below Bottom Coal : Ruiton. 

Alethopteris Sternberg. 
Alethopteris lonchitica Schl., sp. 

1804. Schlotheim, Flora d. Vorwelt, p. 55, pi. xi. fig. 22. 

1820. Filiates lonchiticus, Schlotheim, Petrefactenkunde, p. 411. 

1832 or 1833. Pecopteris lonchitica, Brongt., Hist. d. veyet. foss., p. 275, pi. lxxxiv. figs. 1-7. 

1826. Aletliopteris lonchitidis, Sternb., Vers., vol. i. fasc. iv. p. xxii ; vol. ii, fasc. vii.-viii., p. 142. 

1886. Alethopteris lonchitica, Zeiller, Flore foss. bassin houil. d. Valen., p. 225, pi. xxxi. fig. 1. 
1832 or 1833. Pecopteris urophylla, Brongt., Hist. d. veget. foss., p. 290, pi. lxxxvi. 

Note. — Very common. 
Horizons and Localities. — 

Above Brooch Coal : Himley ; Pensnett ; Shut End ; Oldbury. 

Ten foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley ; 
Cabbage Hall Pit, Netherton. 

Roof of Thick Coal : Tipton ; Ettingshall ; Bradley Colliery, Bilston. 



102 DR ROBERT KIDSTON ON THE 

Roof of Neto Mine Coal: Coseley, near Dudley ; Dibdale, near Gornal ; Clatters- 
hall Colliery, Brettell Lane. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 
Roof of Bottom Coal : Bradley Colliery, Bilston. 
Immediately belotv Bottom Coal : Ruiton. 

Alethopteris decurrens Artis, sp. 

1825. Filicites decurrens, Artis, Antedil. Phyt., pi. xxi. 

1886. Alethopteris decurrens, Zeiller, Flore foss. bassin houil. d. Valen., p. 221, pi. xxxiv. figs. 2, 3 ; 

pi. xxxv. fig. 1 ; pi. xxxvi. figs. 3, 4. 
1832 or 1833. Pecopteris Mantelli, Brongt., Hist. d. veget. foss., p. 278, pi. lxxxiii. figs. 3, 4. 
1834 or 1835. „ „ L. & H., Fossil Flora, vol. ii. pi. cxlv. 

1832. Pecopteris heterqphylla, L. & H., Fossil Flora, vol. i. pi. xxxviii. 

Note. — This species, which is as a rule fairly widely distributed, I have only seen 
from the localities noted below; in the latter mentioned it is not of very rare occurrence. 
Horizons and Localities. — 

Blue Measures, six feet above Brooch Coal: Hamstead Colliery, Great Barr, 

near Birmingham. Collected by Mr H. Insley. 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 

Alethopteris Davreuxi Brongt., sp. 

1832 or 1833. Pecopteris Davreuxi, Brongt., Hist. d. veget. foss., p. 279, pi. lxxxviii. figs. 1, 2. 
1886. Alethopteris Davreuxi, Zeiller, Flore foss. bassin houil. d. Valen., p. 228, pi. xxxii. fig. 1. 
1832 or 1833. Pecopteris Dournaisii, Brongt., ibid., p. 282, pi. lxxxix. fig. 1 (? non fig. 2). 

Note. — Not common. 

Horizon and Localities. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley ; Ettingshall. 

Alethopteris valida Boulay. 

1876. Alethopteris valida, Boulay, Terr, houil. d. Nord de la France, p. 35, pi. i. fig. 8. 

1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 231, pi. xxxiii. figs. 1, 2; 

pi. xxxiv. fig. 1. 
1910. ,, ,, Gothan, in Totonie, Abbild. u. Beschreib. foss. Pflanzen, Lief, vii., No. 125, 

figs. l-4a, b. 
1912. ,, ,, Franke, Beitr. z. Kennt. d. paldoz. Arten v. Alethopteris u. Gallipteridium, 

p. 57, figs. l-4a, b. 
1854. Alethopteris Serli, Geinitz (non Brongt.), Darst. d. Flora d. Hainichen Ebersdorfer Kohlenbassins, 
p. 44, pi. xiv. figs. 3, 4 (1 fig. 5). 

Note. — Very rare. 
Horizons and Localities. — 

Blue Measures above Brooch Coal : Jubilee Pit, Sandwell Park, West 
Bromwich. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 103 



Alethopteris Serli Brongt., sp. 

1832 or 1833. Pecopteris Serlii, Brongt., Hist. d. veget.foss., p. 292, pi. lxxxv. 
1837. ,, ,, L. & H., Fossil Flora, vol. iii. pi. ccii. 

1886. Alethopteris Serli, Zeiller, Flore foss. bassin houil. d. Valen., p. 234, pi. xxxvi. figs. 1, 2 ; 
pi. xxxvii. figs. 1, 2. 

Note. — Very rare. 
Horizons and Localities. — 

Roof of Brooch Coal: Himley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 



Alethopteris Grandini Brongt. 

1832 or 1833. Pecopteris Grandini Brongt., Hist. d. veget.foss., p. 286, pi. xci. figs. 1-4. 
1886. Alethopteris Grandini, Zeiller, Flore foss. bassin houil. d. Valen., p. 227, pi. xxxviii. figs. 1, 2. 
1888. ,, ,, Zeiller, Flore foss. terr. houil. d. Commentry, prem. part, p. 203, pi. xxi. 

figs. 1-8. 

Note. — Very rare. 

Horizons and Localities. — . ■ 

Blue Measures above Brooch Coed: Jubilee Pit, Sandwell Park, West 
Bromwich. 

Roof of Brooch Coal : Holly Hall, near Dudley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 



Alethopteris integra Gothan, sp. 

1906. Desmopteris integra, Gothan, in Potonie, Abbild. u. Besehreib. foss. Pflanzen, No. 64, plate, and 
text-fig. 1. 

Description. — Frond probably large, tripinnate or possibly quadripinnate on lower 
primary pinnse ; rachis striated ; ultimate pinnae broadly linear-lanceolate, opposite 
or alternate, gradually contracting from the base upwards and terminating in a more 
or less blunt point ; uppermost almost entire or bearing broad shallow blunt lobes ; 
lower, more or less deeply divided into pinnules. Pinnules opposite, oblong, decurrent, 
blunt, the basal pinnules united to each other for about ^ to \ of their length ; the 
upper pinnules become more and more united to each other until they assume the 
form of blunt lobes at the base of the oblong obtuse terminal pinnule. Nervation : 
central vein equidistant from margins, decurrent, flexuous, dividing into several 
branches before reaching the apex ; lateral veins alternate, slightly flexuous, dividing 
1-3 times, rather distant ; accessory simple or once-divided veinlets enter the 
pinnules from the rachis and extend into the lateral united portions of the pinnules. 
(Text-fig. 3.) 



104 



DR ROBERT KIDSTON ON THE 



Remarks. — This species has some affinity with Alethopteris Grandini Brongt., sp., 
but differs in its smaller pinnules, which are also more united amongst themselves. It 





.->— ^C- 



•¥' 



:^i 






Text- FIG. 3. — Alethopteris Integra Gothan, sp. Natural size. 




Text-fig. 4. — Alethopteris integra Gothan, sp. Pinnule enlarged 10 times to show the nervation. 



further differs from that species in the central vein of the pinnules, as well as in the 
lateral veinlets being somewhat flexuous and more distant. (Text-fig. 4.) 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 105 

In Alethopteris Grandini, on the other hand, the veins are straight or only slightly 
curved. The general appearance of the two species is also dissimilar ; the more united 
pinnules of Alethopteris Integra impart a much denser character to the frond. 

With Alethopteris valida Boulay it has also a slight similarity, but from that it is 
distinguished by its smaller and very obtuse pinnules as well as by its somewhat 
flexuous veins. The general appearance of the two plants is also essentially distinct. 

Alethopteris Integra, Gothan is apparently a rare species, and the only British 
example I have seen is that given here at text-fig. 3, natural size, and of which a pinnule 
is enlarged 10 times at text-fig. 4. Dr Gothan, who has seen the fossil, agrees with 
me in its identification with his species. 

The specimen was collected by Mr H. W. Hughes, F.G.S., to whom my thanks are 
due for the kind permission to describe and figure it here. 

Horizon and Locality. — Roof of New Mine Coal : Doulton's Clay Pit, Netherton, 
near Dudley. 

Lonchopteris Brongniart. 
Lonchopteris rugosa. 

1835. Lonchopteris rugosa, Brongt., Hist. d. veget. foss., p. 368, pi. cxxxi. fig. 1. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 244, pi. xxxix. figs. 2, 3 ; 

pi. 1. figs. 3, 4. 

Note. — Rare. 

Horizons and Localities. — 

Roof of Brooch Coal: Russell's Hall Colliery, Dudley. Collected by the late 

Sir J. Lundy. 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Old Mine Fireclay = Roof of Stinking Coal : Docking-iron Pit, Delph, Brierley 
Hill. 

Odontopteris Brongniart. 

Odontopteris alpina Sternb., sp. 

1833. Neuropteris alpina, Sternb., Essai Jiore monde prim., vol. ii. fasc. v.-vi. p. 75, pi. xxii. fig. 2. 

1855. Odontopteris alpina, Geinitz, Vers. d. Steinkf. in Sachsen, p. 20, pi. xxvi. fig. 12, pi. xxvii. fig. 1. 
1901. ,, ,, Kidston, Proc. Yorks. Qeol. and Polytech. Soc, vol. xiv. part ii. p. 196, 

pi. xxviii. figs. 1 and la. 
1904. „ ,, Potonie (pars), Abbild. u. Beschreib./oss. Pflanzen, No. 22, fig. 1. 

1870. Xenopteris alpina, Weiss, Zeitsch. d. deut. geol. Gesell., vol. xxii. p. 869. 

1843. Neuropteris confiuens, Gutbier, in Geinitz and Gutbier, Geea v. Sachsen, p. 79. 

Remarks. — The South Staffordshire specimen is similar to that from Yorkshire which 
has been figured by me under the name of Odontopteris alpina, and agrees especially with 
the figure given by Geinitz ; only, my examples are a little smaller in all their parts. 
There is, 1 think, room for doubt as to the specific distinction between Odontopteris 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 14 



106 DR ROBERT KIDSTON ON THE 

alpina and Odontopteris britannica Gutbier,* with which latter species my specimens 



agree more in size. 



At present it seems impossible to settle this point, but, as I have figured f the 
British species under discussion, palaeobotanists will easily recognise the plant which 
I have referred to Odontopteris alpina, and which species is of older date than 
Odontopteris britannica, Gutbier, sp., with which it may possibly be synonymous. 

Potonie unites the Odontopteris genuina Grand' EuryJ and the Odontopteris 
nervosa and Odontopteris densifolia Fortaine and White § with Odontopteris alpina ; 
but these are certainly not the British plant, nor do they appear to be similar to the 
species he himself gives at fig. 1, under the name of Odontopteris alpina. 

Horizon and Locality. — Ten -foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

Neuropteris Brongniart. 

Neuropteris heterophylla Brongt. 
PI. VII. figs. 3, 4. 

1822. Filicites {Neuropteris) heterophyllus, Brongt., Class, d. veget. foss., p. 33, pi. ii. figs. 6a and Qb. 

1830. Neuropteris heterophylla, Brongt., Hist. d. veget. foss., p. 243, pi. lxxi. and pi. lxxii. fig. 2. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 261, pi. xliii. figs. 1, 2; 

pi. xliv. fig. 1. 
1904. „ ,, Kidston, Phil. Trans., ser. B, vol. cxvii. p. 1, pi. i. figs. 1-9 ; text-fig. 1, 

p. 3. 
1911. „ ,, Kidston, Mem. Museeroy. d'hist. nat. d. Belgique, vol. iv. p. 75, text- 

figs. 8, 9 (p. 72), fig. 10 (p. 73), fig. 11 (p. 74). 
1830. Neuropteris Loshii, Brongt., Hist. d. veget. foss., p. 242, pi. lxxii. fig. 1, pi. lxxiii. 
1869. ,, ,, Roehl, Flora d. Steink. Westph., p. 37, pi. xvii. 

1830. Cyclopteris trichomanoides, Brongt., Hist. d. veget. foss., p. 217, pi. lxi. bis, fig. 4. 

Remarks. — Since preparing the original description of the fructification of 
Neuropteris heterophylla, \\ some additional specimens have been collected by Mr 
H. W. Hughes, and two of these are described here, as they are more perfect than 
those first discovered. IF 

That shown on PI. VII. fig. 3 has probably reached maturity. It is narrow oval, 
with a somewhat flattened base by which it is attached to the expanded apex of the 
rachis, from whose side springs a pinnule showing the typical form and nervation 
of the species. The cupular structure at the base of the seed is not so well seen 
in this example as in some of those previously figured, and especially those at figs. 
5 to 8 of the original paper. 

In the specimen here figured the stalk terminates in a solid conical expansion, 

* Gutbier, A bdr. u. Verstein. d. Zwick. Schwarz, p. 68, pi. ix. figs. 8-11 ; see also Gkinitz, I.e., p. 21, pi. xxvi. 
figs. 8, 9. t hoc. cit. 

% Flore carbon, du depart, de la Loire, p. 115, 1877. Zeiller, Flore foss. terr. houil. d. Commentry, p. 219, pi. xxiv. 
figs. 1, 3 ; pi. xxv. figs. 1, 2 ; pi. xxxi. fig. 1. 

§ Fortaine and White, Permian Flora, pp. 52 and 54, pi. x. figs. 1-3, 1880. || Phil. Trans., I.e., p. 1. 

1 These two specimens are figured in Kidston, Mem. Musee roy., I.e., text-figs. 8, 9. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 



107 



which is well seen in the figure, and to which the seed is attached. The other half 
of the nodule shows a small foliar fragment which might be the remains of the cupular 
leaf that springs from the apparent margin of this basal cushion, as seen in figs. 6 
and 8 of the original paper. 

The seed, which is 5 cm. long and 2 cm. broad, gradually contracts into a micropylar 
beak, which is about 075 cm. long but may not be quite complete. 

The other example figured here on PL VII. fig. 4 is an immature individual 
2 - 02 cm. long and 0*80 cm. broad. The upper part of the seed is prolonged into a 




Text-fig. 5. — Neuropteris heterophylla Brongt. Immature seed, enlarged 3 times. 

blunt micropylar beak, and the base is attached to a thickening very similar to that 
described above. To the side of the cushion which bears the seed is attached a foliage 
pinnule of the ordinary type. The specimen is seen enlarged 3 times at text-fig. 5. 
Neuropteris heterophylla is very common and widely distributed on the coal field. 
Horizons and Localities. — 

Roof of Brooch Coal: Himley ; Bilston ; Holly Hall, near Dudley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley ; 

Tipton ; Ettingshall ; Cabbage Hall Pit ; Doulton's Clay Pit, Netherton. 
Roof of Thick Coal : Bradley Colliery, Bilston. 
" Whitestone" : Race Course Pit, Round Oak. 
Roof of Stinking Coal : Clattershall Colliery, Brettell Lane. 
Roof of New Mine: Dibdale, near Gornal ; Doulton's Clay Pit, Netherton. 
Roof of Fireclay Coal : Doulton's Clay Pit, Netherton. 
Immediately below Bottom Coal : Ruiton, near Sedgley. 



108 



DR ROBERT KIDSTON ON THE 



Neuropteris tenuifolia Schl, sp. 

1820. Filicites tenuifolius, Schlotheim, Petrefactenlcunde, p. 405, pi. xxii. fig. 1. 

1830. Neuropteris tenuifolia, Brongt., Hist. d. veget.foss., p. 241, pi. lxxii. fig. 3. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 273, pi. xlvi. fig. 1. 

Note. — This species seems to be rarer in the South Staffordshire Coal Field than 
one would naturally expect. This apparent rareness may be the accident of collecting. 
Horizons and Localities. — 

Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr. 

near Birmingham. Collected by Mr F. Stephens. 
Roof of Brooch Coal: Tividale. 



Neuropteris gigantea Sternb. 
PI. VI. figs. 1 to 7. 

1823. Osmunda gigantea, Sternb., Essai /lore monde prim., vol. i. fasc. ii. pp. 32, 37, pi. xxii. 

1830. Neuropteris gigantea, Brongt., Hist. d. veget.foss., p. 240, pi. lxix. 

1832. „ „ L. & H., Fossil Flora, vol. i. pi. Hi. 

1886. ,, „ Zeiller, Flore foss. bassin houil. d. Valen., p. 258, pi. xlii. fig. 1. 

1892. „ ,, Potonie, "Ueber einige Carbonfarne," iii. Theil, p. 22, text-figs. 1-4; pi. ii. 

figs. 1, 2; pi. iii. figs. 1-4; pi. iv. figs. 1, 2 (Jalirb. d. k. preuss. geol. 

Landesanstalt fur 1891). 
1910. „ ,, Renier, Paleont. d. terr. houil., pi. c. 

1892. Neuropteris Zeilleri, Potonie, ibid., pp. 22, 32, fig. 5. 
1899. Neuropteris pseudogigantea, Potonie^ Lehrb. d. Pflanzenpal., p. 113, fig. 102. 

Remarks. — Dr Potonie has divided the plants which formerly were placed under 
Neuropteris gigantea Sternb. into two species, Neuropteris gigantea Sternberg and 
Neuropteris pseudogigantea (= Neuropteris Zeilleri Potonie). The difference between 
these two " species" he has given in tabular form, which I summarise here. # 



Neuropteris gigantea Sternb. 

Pinnules. 
1. Sickle-shaped, bent. 

2 Obliquely elongate, ovate cordate, narrowing 
towards the apex. 

3. In general longer than in Neuropteris Zeilleri. 

4. Breadth (measured in the middle) in proportion 

to the length as 1 to 3 or more. 

5. Middle nerve not observable. From below 

dividing into veinlets. At the most only a 
feeble indication of a middle vein at the base of 
the pinnule ; generally smooth without furrow ; 
sometimes a furrow is weakly indicated. 

6. Anastomoses between the veins not at all 

frequent. 



Neuropteris Zeilleri Potonie (subsequently named 
by Potonij6 Neuropteris pseudogigantea (I.e.)). 

Pinnules. 
Quite straight (im ganzen gerade). 
Broadly linear, margins parallel. 



3. Length to about 25 millimetres. 

4. Breadth in regard to the length in rjroportion of 

1 to 21 or 2£. 

5. Middle nerve extending to above the centre of 

the pinnule, distinctly seen and marked by a 
clear furrow ; mid-vein breaks up into veinlets 
about the middle of the pinnule. 

6. Anastomoses between the veins apparently more 

frequent. 



* Potoni£, I.e., p. 31. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 109 

If these characters were constant, then there might be ground for the separation 
of Neuropteris gigantea Sternb. into the proposed two species ; but unfortunately they 
appear to be very inconstant. 

Between the sickle-shaped and " straight " pinnules there is every graduation 
of form. In fact, the pinnules of Neuropteris gigantea, to use the old name 
collectively, are extremely variable. Almost invariably the pinnules are slightly 
oblique — that is, one side is slightly convex and the other concave. This is seen in 
the figure given by Zeiller,* which is the type of Neuropteris Zeilleri ( = Neuropteris 
pseudogigantea) Potonie. In fact, a pinnule of "Neuropteris gigantea" with straight 
sides is very seldom seen. 

The pinnules of Neuropteris gigantea, like others of the genus, are articulated 
to the rachis, and, like our Osmunda regalis, seem to have been shed when the fronds 
ripened to decay. Hence it is comparatively seldom that one gets large specimens 
of this species showing the pinnules attached to the rachis. The much more common 
mode of occurrence is to find the isolated pinnules scattered over the surface of the 
shales containing them. 

Among these isolated pinnules it is easy to find connecting forms from Neuropteris 
gigantea Sternb. to Neuropteris pseudogigantea Potonie. As far as the form of the 
pinnule is concerned, it seems impossible on this character to separate the " species." 
Even on the type-figure of Sternberg t some of the pinnules are falcate or sickle- 
shaped while others are straight, and a few "straight" pinnules also occur on the 
specimen figured by Potonie himself. J Accordingly, if you removed the pinnules from 
the rachis of these two specimens and scatter them on a slab, you would find both 
species represented. As to size, the pinnules vary much even on the same specimen. 

In regard to the nervation, neither is this constant in the two forms, for I possess 
a British specimen (No. 4519) which is typical in all respects with Neuropteris gigantea 
Sternb. (Potonie), except that the mid-vein extends well up the pinnule and does 
not break up into veinlets at or near the base. I have also a specimen from Prussia 
showing the same character. An anastomosing of the veins I have never observed 
to occur in any form of Neuropteris gigantea. 

I have therefore failed to satisfy myself as to the specific value of Neuropteris 
pseudogigantea Potonie, and therefore include it with Neuropteris gigantea Sternberg. 

Some specimens of Neuropteris gigantea from the South Staffordshire Coal Field 
are given on PI. VI. figs. 1 to 7, and these illustrate the form almost invariably found 
there. All the examples shown on the Plate are preserved in ironstone nodules, and 
show the great beauty with which the details of structure can be preserved in 
this matrix. 

At fig. 1 a fragment of an ultimate pinna is given. This specimen would probably 
fall to be placed under the Neuropteris pseudogigantea Potonie by those who uphold 

* Flore f oss. bassin houil. d. Valen., pi. xlii. fig. 1 ; well seen in fig la. 
t PI. xxii. t PI. iii. 



110 DR ROBERT KIDSTON ON THE 

that species. The pinnules are slightly oblique, and a single pinnule of a similar form 
is given at fig. 2, enlarged 2^- times to show the nervation. The central vein can 
be traced upwards from the base for about ^ the length of the pinnule, where it 
breaks up into diverging veinlets. This figure, as well as fig. 3, shows the pinnae 
terminating in two little diverging pinnules, not in a single terminal pinnule as in 
almost all the other members of the genus At fig. 4 the rachis of the pinna is seen 
to bifurcate ; and I have another small specimen from Elsecar Colliery, Wentworth, 
Yorkshire, which probably comes from the Barnsley Bed, Westphalian Series, which 
shows a double bifurcation of the rachis (K. 4019). 

Some Spiropte?^is conditions of the plant are given at figs. 5 to 7. That at fig. 5 
shows several pinnse partially inrolled, and much larger specimens in this state occur 
in the ironstone nodules of Coseley : one in my collection, No. 4011, is 18 cm. long, 
and shows many pinnse partially developed. Less advanced stages of development 
are seen at figs. 6 and 7, all of which are sufficiently well preserved to admit of 
a satisfactory determination of the species. The most interesting point in the specimens 
is the presence of the thick covering of scales on the rachis, which must have been 
very fugaceous and slightly attached, as one almost never sees any cicatrices to 
indicate their former presence on the rachides of the developed pinnse, though they are 
clearly shown on the specimen figured by Potonie on his pi. iii. # 

Neuropteris gigantea Sternb. is very common in the South Staffordshire Coal Field. 
Horizons and Localities. — 

Blue Measures, above Brooch Coal : Jubilee Pit, Sandwell Park, West Bromwich. 

Roof of Brooch Coal : Shut End ; Bradley Colliery, Bilston ; Holly Hall, 
near Dudley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley ; 
Moxley, near Bilston. 

Immediately below Thick Coal : Yew Tree Colliery, Rowley Regis. 

" Old Clay" = Roof of Stinking Coal : The Delf, Brierley Hill. 

" Whitestone " : Race Course Pit, Round Oak. 

Roof of New Mine: Merryhill Colliery, Mount Pleasant, Brierley Hill; Dibdale 
Colliery, near Gornal. 
• Roof of Fireclay Coal: Doulton's Clay Pit, Netherton. 

Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 

Roof of Bottom Coal: No. 120 Pit, Coneygre Colliery, Tipton. 

Immediately below Bottom Coal: Ruiton, near Sedgley. 

Neuropteris rarinervis Bunbury. 

1847. Neuropteris rarinervis, Bunbury, Quart. Journ. Geol. Hoc, vol. iii. p. 425, pi. xxii. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 268, pi. xlv. figs. 1-4. 

1870. Neuropteris coriacea, Lesqx., Rept. Geol. Survey of Illin., vol. iv. p. 387, pi. viii. figs. 7, 8. 

* hoc. cit. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. Ill 

Note. — Rare. 

Horizons and Localities. — 

Shale, thirty yards above Thick Coal : Hamstead Colliery, Great Barr, near 
Birmingham. 

Immediately below Bottom Coal : Ruiton, near Sedgley. 

Neuropteris Grangeri Brongt. 

1830. Neuropteris Grangeri, Brongt., Hist. d. veget. foss., vol. i. p. 237, pi. lxviii. fig. 1. 
1879. „ ., Lesqx., Coal Flora, p. 105, pi. xiii. fig. 9. 

' 1890. ,, „ Zeiller, Flore foss. d. bassin houil. et perm. d'Autun et d'fipinac, p. 145, 

pi. xi. fig. 6. 

Note. — Very rare. 
Horizons and Localities. — 

Roof of Tliick Coal: Bradley Colliery, Bilston. 

Roof of Fireclay Coal: Doulton's Clay Pit, Netherton. 

Neuropteris Schlehani Stur. 

1877. Neuropteris Schlehani, Stur, Culm Flora, Heft ii., p. 289, pi. xxviii. figs. 7, 8a, b, c. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 280, pi. xlvi. fig. 3 ; pi. xlvii. 

figs. 1, 2. 
1879. Neuropteris Elrodi, Lesqx., Coal Flora, vol. i. p. 107, pi. xiii. fig. 4. 
1877. Neuropteris Dluhoschi, Stur, ibid., p. 289, pi. xxviii. fig. 9. 

Note. — This species, which is so common in some of the Continental coal fields of 
Westphalian age, is undoubtedly a very much rarer species in Britain. 
Horizons and Localities. — 

Brooch Coal Binds : Oldbury. 

Shale above Thick Coal : Dudley. 

Roof of Bottom Coal: Tipton. (British Museum, No. 52,775 and v. 1301.) 

Neuropteris obliqua Brongt., sp. 

1832. Pecopteris obliqua, Brongt., Hist. d. veget. foss., p. 320, pi. xcvi. hgs. 1-4. 
1886. Neuropteris obliqua, Zeiller, Flore foss. bassin houil. d. Valen., p. 284, pi. xlviii. figs. 1-7. 
1911. ,, ,, Kidston and Jongmans, " Sur la fructification de Neuropteris obliqua 

Brongt.," Archives Neerland. d. Sciences exactes et nat., ser. iii., B, vol. i. 
p. 25, pi. 
1903. Neurodontopteris impar, Weiss, MS., in Potonie, " Uber einige Carbonfarne," iv. Tlieil, p. 1, 

pi. i. (Jahrb. d. kbnigl. preuss. geol. Landesanst. fur 1902). 
1836. Neuropteris heterophylla, L. & H. (non Brongt.), Fossil Flora, vol. iii. pi. clxxxiii. 
1886. Neuropteris acuminata, Zeiller (non Schloth.), Flore foss. bassin houil. d. Valen., p. 255, pi. xli. 

fig. 4. 
1911. Neuropteris impar, Kidston, Mem. Musie roy. d'hist. nat. de Belgique, vol. iv. p. 83, pi. viii. 

figs. 1, la, 2, 3, 3a. 



112 DR ROBERT KIDSTON ON THE 

Remarks. — A polymorphic species, of which the seed is very similar in size and 
external characters to that of Neuropteris heterophylla Brongt. It would, in fact, be 
very difficult, if even possible, to allocate to their respective species the seeds of these 
two Pteridosperms, if separated from branches bearing their distinctive foliage. 
Both the obliqua and impar form of the plant occur in this coal field. 
Horizons and Localities. — 

Blue Measures, six feet above . Brooch Coal : Hamstead Colliery, Great Barr, 

near Birmingham. Collected by Mr H. Insley. 
Brooch Coal Binds : Oldbury. 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 

Neuropteris Osmundse Artis, sp. 

1824. Filicites Osmundx, Artis, Antedil. Phyt., p. 7, pi. vii. 

1829. Neuropteris auriculata, Brongt., Hist. d. veget. foss., vol. i. p. 236, pi. lxvi. 
1844. ,, ,, Germar, Vers. d. Steink. v. Wettin u. Lbbejun, p. 9, pi. iv. 

1893. Neurodontojpteris auriculata, Potoni^, Flora d. Roihl. v. Thuringen, p. 124, pi. xvi. figs. 1, 2. 

1830. Neuropteris Villiersii, Brongt., Hist. d. veget. foss., p. 233, pi. lxiv. fig. 1. 

Note. — Not common. 

Horizons and Localities. — Ten-foot Ironstone Measures : Tipton ; Clayscroft 
Openwork, near Dudley ; Ettingshall. 

Neuropteris Scheuchzeri Hoffman. 

1826. Neuropteris Scheuchzeri, Hoffm., in Keferstein's Teuchland geognostisch dargestellt, vol. iv. 

p. 156, pi. i. b figs. 1-4. 
1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 251, pi. xli. figs. 1-3. 

1888. ,, ,, Kidston, Trans. Roy. Soc. Edin., vol. xxxiii. p. 356, pi. xxiii. figs. 1, 2. 

1830. Neuropteris angustifolia, Brongt., Hist. d. veget. foss., p. 231, pi. lxiv. figs. 3, 4. 
1832. Neuropteris cordata, L. & H. (non Brongt.), Fossil Flora, vol. i. pi. xli. 
1858. Neuropteris hirsuta, Lesqx., in Rogers, Geol. of Pennsyl., vol. ii. p. 857, pi. iii. fig. 6, pi. iv. 

figs. 1-16. 
1879. „ „ Lesqx., Goal Flora, p. 89, pi. viii. figs. 1, 4, 5, 7, 9, 12. 

Note. — Not common. 
Horizons and Localities. — 

Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr. 
near Birmingham. Collected by Mr H. Insley. 

Roof of Brooch Coal : Himley. 

Immediately below Bottom Coal : Ruiton, near Sedgley. 

Neuropteris Carpentieri Kidston, n. sp. 
PI. VIII. figs. 1-7. 

1911. Potoniea aff. adiantiformis, Carpentier (non Zeiller), " Sur quelques fructifications et inflorescences 
du Westphalien du Nord de la France," p. 13, pi. xvi. figs. 2, 3 (Revue generate de hot., 
vol. xxiii.). 

1911. Potoniea adiantiformis, Carpentier (non Zeiller), ibid., p. 13, pi. xvi. fig. 1 (fig. 21). 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 113 

Description. — Frond probably of large size. Ultimate divisions of frond with thick, 
strong, striated rachis, each of which terminates in a sub-cyclopteroid pinnule of thick 
substance whose ventral surface shows, when the upper layer bearing the sporangia has 
been removed, the entrance of several strong veins from the rachis into the pinnule, 
which radiate from the base and dichotomise two to three times in their course to the 
margin. Upper or adaxial surface of pinnule bearing densely packed, narrow, elongated 
microsporangia about 4 mm. long and about 0*5 mm. broad. Microspores small, 
round or somewhat oval, averaging from p. 45 to m 60 in diameter. Sterile pinnule 
neuropteroid, terminating a short thick rachis, from which two veins seem to enter the 
pinnule that give off dichotomously divided lateral veinlets, and in the upper part break 
up into several branchlets. 

Remarks. — Several specimens of this fossil are given on PI. VIII. That seen at 
fig. 4 is the largest specimen met with, but the rachis is only seen at the upper part. 
Here it is 2 '5 mm. wide immediately below its attachment to the pinnule. The upper 
part of this specimen is enlarged 2 times at fig. 4a, to show the striated rachis caused 
by numerous Myelopteris-Yike veins which pass up it to enter the cyclopteroid pinnules. 
This example shows the ventral aspect of the pinnule, from which the sporangia have 
been removed, along with the layer to which they were attached, by adherence to the 
other half of the nodule ; and a similar condition occurs on the other pinnules seen on 
this specimen. In almost all cases the fracture of the stone passes underneath the 
layer bearing the sporangia, which adhere to the counterpart of the fossil and there 
exhibit the under surface of the layer to which they are attached. 

At fig. 1 the pinnule is bent up and also shows its ventral aspect. This figure 
might give the idea that this pinnule was peltate, but the petiole joins on to the basal 
margin of the pinnule, whose more distal portion is curiously bent up and partially 
buried in the matrix. 

Fig. 2 shows natural size the ventral view of two pinnules which terminate a 
dichotomy ; that at a is incomplete, the upper portion being broken off, while that at b 
probably shows the complete pinnule, or only a small part of the upper margin is 
missing. Another pinnule is given at fig. 6, enlarged 2 times. This also terminates 
one branch of a dichotomy, but only a small portion of the corresponding pinnule is 
visible. This, I am inclined to think, is a sterile pinnule, not one whose ventral surface 
has become exposed by the removal of the sporangial layer, and illustrates one of those 
intermediate forms between the sterile and microsporangiate cyclopteroid pinnules. 
A further stage of transition from the cyclopteroid to the normal Neuropteroid pinnule 
is seen on the specimen shown at fig. 7, where the fossil is given natural size, but the 
details of the structure are better seen at fig. 7a, which is enlarged 2 times. Here 
the pinnule lettered a is that seen at a, fig. 7. The upper portion of this pinnule is 
unfortunately broken over, but the portion preserved is typical of an ordinary sterile 
pinnule of Neuropteris. Two veins appear to enter it, but this is no very unusual 
character, as normally more than one vein enters the pinnules of Neuropteris ovata 
TRANS. ROY. SOC. ED1N., VOL. L. PART I. (NO. 5). 15 



114 DR ROBERT KIDSTON ON THE 

and Neuropteris obliqua and probably other species ; but this condition can only be 
observed in favourably preserved specimens. The evidence, then, for referring these 
specimens to Neuropteris seems to be fairly conclusive. 

Let us now turn to the specimen given at figs. 5a and 56. These show the two 
halves of the same nodule, enlarged 2 times. At fig. 5 a we are looking at the upper 
surface of the pinnule, from which the cushion on which the sporangia sat has been torn 
off. The under surface of this cushion is seen at a, fig. 56 ; and the part lettered a! on 
the same figure is a piece of the pinnule broken off a, fig. 5a, and is not sporangial. 
At 6 and c, fig. 56, parts of the pinnule also are seen. On no part of this specimen are 
any sporangia visible, as far as I can observe. 

The only specimen which clearly shows the sporangia is that seen at figs. 3a and 36, 
of which both halves of the nodule are given, enlarged 2 times. Here we are looking 
down upon the upper surface of the sporangial disc. At fig. 3a they are much flattened 
and displaced, but at a on fig. 36 they are seen distinctly extending past the margin of 
the disc. The greater part of the surface of this specimen exhibits small hummocky 
roughnesses. These, I believe, are the points from which sporangia have been removed, 
and which now adhere in a crushed condition to the surface of the other half, fig. 3a. 

Some of the specimens, when split open, contained some portions of the plant in a 
carbonaceous condition adhering to the impression. Some of this material was treated 
by the maceration process, and numerous spores were obtained from both the specimens 
so examined, some of which are seen at PL VIII. fig. 8. This clearly proved the 
sporangial nature of the bodies seen on the specimen given at figs. 36 and 3a. The 
spores show a triradiate ridge, fig. 8a. 

That the fossils just described are the microsporangial organs of a Neuropteris can 
scarcely be doubted, but it is impossible to determine the species to which they belong ; 
for though at a on fig. 7a there is clearly a Neuropteroid pinnule, it is so closely 
associated with the fertile pinuules that it probably does not represent the true form 
and arrangement of the veins in the sterile pinnules. Several species of Neuropteris 
occur at the same locality and on the same horizon, and of these Neuropteris 
heterophylla and Neuropteris gigantea were extremely common. That our specimens 
do not belong to Neuropteris heterophylla is seen from the much more delicate 
structure of the petioles on the probable but imperfectly preserved microsporangial 
specimen I figured some years ago. # 

On the other hand, there is no satisfactory evidence for referring these micro- 
sporangia to Neuropteris gigantea, though pinnules of this species sometimes occur in 
the same nodules, as their presence there is possibly merely an accidental association. 
I therefore apply a new specific designation to these fossils, and have pleasure in 
naming them after M. l'Abbe" Carpentier, who was the first publicly to call the 
attention of botanists to them. In his paper describing them he mentions the original 
MS. name, Cupulina Jilicoides, which I applied to these fossils, but this I suppress for 

* Trans. Roy. Soc. Edin., vol. xxxiii. p. 150, pi. viii. fig. 7, 1887. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 115 

the name given above, as since that provisional name was proposed I now feel satisfied 
that the fossils belong to the genus Neuropteris. 

Of the specimens figured by the Abbe Carpentier, that given on his pi. xvi. fig. 1 
at b, which he refers to Potoniea adiantiformis Zeiller, and those he gives as doubtful 
at figs. 2 and 3, are certainly the same plant as that described here. 

In Potoniea Zeiller * the pinnules are cuneate like those of Adiantum, with delicate 
foot -stalks, not cyclopteroid and attached to a very thick rachis ; and, more important, 
the sporangia in Potoniea seem to be confined to the margins, not covering the whole 
upper surface of the pinnules as in Neuropteris Carpentieri, where the sporangial 
pinnules must have had a very similar appearance to a hair-brush if we turn it upside 
down, the bristles representing the sporangia and the handle the petiole. 

Neuropteris Carpentieri in some states of preservation has a slight resemblance to 
Dictyothalamus Schrollianus Gopp.,t but the apparently distichous arrangement of the 
"buds" on that fossil to a common axis indicates an entirely different class of plant, 
and one which does not appear to have any affinity to the Pteridosperms with which 
undoubtedly Neuropteris Carpentieri has. 

All the figured specimens have been collected by Mr H. W. Hughes, F.G.S. 

Horizon and Localities. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley ; Ettingshall. 

Neuropteris sp. 

Remarks. — Specimens of a Neuropteris have been found at Sandwell by Mr H. W. 
Hughes which have somewhat the form of Neuropteris tenuifolia, but with a much 
wider nervation and the pinnules more distant. 

Horizon and Locality.- — Blue Measures, above Brooch Coal : Jubilee Pit, Sandwell 
Park, West Bromwich. 

Aphlebia Presl 
Aphlebia crispa G-utbier. 

1835. Fucoides crispus, Gutbier, Abdr. u. Verst. d. Zwick. Schivarzkohl, p. 13, pi. i. fig. 11 (? pi. vi. 

fig. 18). 
1886. Aphlebia crispa, Zeiller, Flore foss. bassin houil. d. Valen., p. 304, pi. li. figs. 1, 2. 
1855. Schizopteris lactuca, Geinitz, Vers. d. Steinkf. in Sachsen, p. 19, pi. xxvi. figs. 1. 
1869. „ ,, Roehl, Foss.-Flora d. Steink. Form. Westph., p. 47, pi. xviii. 

1869. Rhacophyllum laduca, Schimper, Traite d. paleont. veget., vol. i. p. 684, (? pi. xlvi. fig. 1), 

pi. xlvii. fig. 1 (non fig. 2). 

Note. — Very rare. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

* "Flore foss. bassin houil. d'Heraclee," p. 52, Mem. Soc. geol. d. France: FaUont. Mem., No. 21, 1899. 
t Perm. Flora, p. 164, pi. xxiv. figs. 4-6, pi. xxv. figs. 1-4. 



116 DE ROBERT KIDSTON ON THE 

Aphlebia sp. 

1835. Fucoides filiciformis, Gutbier (pars), Ver. d. Zwick. Schwarzk., p. 11, pi. i. fig. 6 (wow figs. 3, 6, 
7, 8, 13). 

Remarks. — Under the name of Fucoides filiciformis, Gutbier appears to have 
included more than the remains of one species. That given at his fig. 6, pi. i., seems 
to be quite distinct from those forms usually included under the name of Aphlebia 
filiciformis, and to which perhaps most, if not all, of his other figures belong. 

To this fig. 6 I refer a small specimen contained in an ironstone nodule, which, 
though not too distinctly preserved, shows the slender ramification of this figure ; but 
until better examples are discovered it is sufficient to record it provisionally under 
Aphlebia sp. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

Spiropteris. 

1869. Schimper, Traite d. paleont. veget., vol. i. p. 688. 

Spiropteris sp. 
PI. VI. figs. 8, 9, and 9a. 

Remarks. — This name has been proposed by Schimper, not in the sense of a genus, 
but as a convenient designation under which may be placed circinately coiled fronds of 
ferns or fern-like plants or specimens of fronds that are not fully developed, but which 
may be of themselves interesting, though they cannot be identified with any definite 
species. Such specimens of Spiropteris sometimes show the dichotomy of the petiole 
and the presence of hairs or scales that are produced for the protection of the growing 
parts, but which are usually shed at an early period of the fronds' development, and 
often leave no trace of their former presence, as is usually the case in Neuropteris 
gigantea Sternb., where, as already mentioned, the rachis in the early condition bears a 
dense covering of scales, but which when shed seldom ever leave any scars to attest 
their former presence.* 

Two specimens of Spiropteris are given on PL VI. figs. 8-9. Fig. 9 shows natural 
size a complete young frond with a very thick petiole, bearing transverse scars probably 
left by fallen scales. At the top, the petiole bifurcates into two circinately coiled 
branches which bear the circinately coiled pinnae. The upper part of this fossil is seen 
enlarged 2 times at fig. 9a. Another specimen is given at fig. 8. Here also the 
petiole dichotomises, and is seen to be marked with close transverse scars and divided 
into two arms which in turn bear primary pinnae, and these again secondary pinnae, all 
more or less circinately coiled. 

* See PI. VI. figs, la, b, c. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 117 

That shown at fig. 8 was collected by Mr H. W. Hughes, and that at fig. 9 by 
Mr W. Madeley, and I am indebted to these two friends for the interesting 
specimens. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Equisetacese. 

Equisetites Sternberg. 

Equisetites Hemingway! Kidston, sp. 

1892. Equisetum Hemingwayi, Kidston, Ann. and Mag. Nat. Hist., p. 138, figs, a, b. 

1898. Equisetites Hemingwayi, Seward, Fossil Plants, vol. i. p. 263, fig. 57a. 

1901. „ „ Kidston, Proc. Yorks. Geol. and Polyt. Soc., vol. xiv. p. 198, pi. xxxiv. 

fig. 3. 
1911. ,, ,, Jongmans, Anleit. Karbonpflanzen West-Europas, p. 30, figs. 35 and 37. 

Note. — Very rare. I have only seen a single specimen of Equisetites Hemingwayi 
from this coal field. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Calamites Suckow. 
Calamites Suckowi Brongt. 

1828. Calamites Suckowi, Brongt., Hist. d. veget.foss., p. 124 (pi. xiv. fig. 6?), pi. xv. figs. 1-6 ; pi. xvi. 

_ (fig- 1 ?) 2, 3, 4. 
1876. „ ,, Weiss, Steinkohlen Calamarien, part i. p. 123, pi. xix. fig. 1 ; part ii.. 1884, 

p. 129, pi. ii. fig. 1 ; pi. iii. figs. 2, 3; pi. iv. fig. 1 ; pi. xvii. fig. 5 ; 

pi. xxvii. fig. 3. 
1886. „ „ Zeiller, Flore foss. bassin houil. d. Valen., p. 333, pi. liv. figs. 2, 3; pi. Iv. 

fig. 1. 

Note. — Common. 
Horizons and Localities. — 

Above Brooch Coal: Oldbury ; Himley ; Shut End, near Kingswinford ; Holly 

Hall, near Dudley. 
Shales, thirty yards above Thick Coal : Hamstead Colliery, Great Ban, near 

Birmingham. 
Roof of Thick Coal: Bradley Colliery, Bilston. 
Immediately below Thick Coal : Yew Tree Colliery, Rowley Regis. 
White Ironstone : Sandwell Park Colliery, West Bromwich. 
Roof of Bottom Coal: No. 120 Pit, Coneygre Colliery, Tipton. 
? Parkfield, near Wolverhampton. 

? Russell's Hall, Dudley. 



118 DR ROBERT KIDSTON ON THE 

Calamites Cisti Brongt. 

1828. Calamites Cistii, Brongt., Hist, d. veget.foss., p. 129, pi. xx. 

1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 342, pi. lvi. figs. 1, 2. 

Note. — Not very common. 

Horizon and Localities. — Roof of Brooch Coal : Oldbury, near Dudley ; Shut 
End, near Kingswinford ; Pensnett. 

Calamites undulatus Sternb. 

1826. Calamites undulatus, Sternb., Essai flore monde prim., vol. i. fasc. iv. p. 26; vol. ii. fasc. v.-vi. 

p. 47, pi. i. fig. 2 (? pi. xx. fig. 8). 
1828. ,, ,, Brongt., Hist. d. veget.foss., p. 127, pi. xvii. figs. 1-4. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 338, pi. liv. figs. 1 and 4. 

Note. — This plant, usually so common, I have only received from a single locality. 
This apparent rareness of Calamites undulatus in the South Stafford Coal Field may 
possibly arise from the plant having been passed over in collecting. 

Horizon and Locality. — Roof of Brooch Coal: Shut End, near Kingswinford. 

Calamites Waldenburgensis Kidston. 

1828. Calamites approximatus, Brongt. (non Schl.) (pars), Hist. d. veget.foss., p. 133, pi. xxiv. figs. 2, 3 

figs. 4, 5). 
1855. ,, ,, Geinitz (pars), Vers. d. Steinkf. in Sachsen, p. 7, pi. xi. fig. 5, pi. xii. 

fig. 3. 

1887. ,, ,, Stur (pars), Calamarien d. Carbon-Flora d. Schatz. Schichten, p. 119, 

pi. v. fig. 3, pi. viii. fig. 4. 
1893. Calamitina approximata, Kidston, Trans. Roy. Soc. Edin., vol. xxxvii. p. 311, pi. ii. figs. 5, 6. 
1903. Calamites Waldenburgensis, Kidston, Trans. Roy. Soc. Edin., vol. xl. p. 789. 
1911. ,, ,, Jongmans, Anleitung, vol. i. p. 57, figs. 65, 66. 

Note. — Rare. 

Horizons and Localities. — 

Ten-foot Ironstone Measures : Cabbage Hall Pit, Netherton. 

Roof of Fireclay Coal : Doulton's Clay Pit, Netherton, near Dudley. 

Below Bottom Coal : Ruiton, near Sedgley. 

Calamites Schutzei Stur. 

1887. Calamites Schutzei, Stur, Calamarien d. Carbon-Flora d. Schatz. Schichten, p. 131, pi. iii. figs. 2, 
26 ; pi. iv. ; pi. iv./,, fig. 1 ; pi. xvii. fig. 2 ; text-figs. 33-38. 

Note. — Rare. 

Horizon and Locality. — Roof of Fireclay Coal : Doulton's Clay Pit, Netherton, 
near Dudley. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 119 

Calamites ramosus Artis. 

1825. Calamites ramosus, Artis, Antedil. Phyt., pi. ii. 

Note. — Frequent. 
Horizons and Localities. — 

Roof of Brooch Coal : Oldbury ; Shut End, near Kingswinford. 

Brooch Binds Ironstone : Pensnett ; Himley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 

Roof of Fireclay Coal : Doulton's Clay Pit, Netherton, near Dudley. 

Below Bottom Coal : Ruiton, near Sedgley. 

Calamites Britannicus Weiss. 

1888. Eucalamites {Calamites) Britannicus, Weiss, Ann. and Mag. Nat. Hist., ser. 6, vol. ii. p. 131, 

pi. vii. 
1911. Calamites Britannicus, Jongmans, Anleitung, p. 123, fig. 118. 

Note. — Very rare. Only a single specimen of this species has been found by 
Mr C. Beale in 1888. 

Geinitz unites this species with his Calamitina oculata, but I believe the two 
species are quite distinct.* 

Horizon and Locality. — Above Thick Coal: Shut End near Kingswinford. 

Calamites sp. : Rhizomatic tuber. 
PL IX. fig. 2. 

Remarks. — A rhizomatic tuber of -a Catamite is shown on PL IX. fig. 2, natural size. 
These tuberous structures, though well known to occur on some species of Equisetum, 
have not been previously known to occur on Calamites, as far as I am aware. The 
specimen, which is preserved in an ironstone nodule, shows an inflation of the rhizome 
about 2 '50 cm. long and 1 cm. wide at its centre. The tuber consists of three 
internodes, measuring respectively 1'30 cm., 0*70 cm., and 0'50 cm., of which the 
centre internode is the widest ; the two end ones narrow into the rhizome, which, 
where they join it, is about 3 mm. wide. At one end of the fossil only one internode 
of the rhizome is preserved, which is apparently incomplete, but measures 1 cm. in 
length. At the other end the rhizome decreases quickly in width and is much 
thinner, and contains seven very short internodes. Both the tuber and rhizome are 
distinctly ribbed, and rootlets are given off from all the nodes. 

This interesting specimen was collected by Mr H. W. Hughes. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

* Geinitz, " Die Calainarien der Steinkoblen-Formation u. d. Roethl. in Dresdener Museum," Mitt. Jcongl. Miner.- 
Geol. u. Praehis. Museum im Dresden, Vierzehntes Heft, p. 12, pi. i. figs. 1, 2, 1898. 



120 DR ROBERT KIDSTON ON THE 

Asterophyllites Brongniart. 
Asterophyllites equisetiformis Sohl., sp. 

1804. Schlotheim, Flora d. Vorwelt, p. 30, pi. i. figs. 1, 2; pi. ii. fig. 3. 

1820. Casuarinites equisetiformis, Schloth., Petrefactenkunde, p. 397. 

1828. Asterophyllites equisetiformis, Brongt., Prodrome, p. 158. 

1845. ,, ,, Gerniar, Vers. v. Wettin u. Lbbejun, p. 21, pi. viii. 

1886. ,, „ Zeiller, Flore foss. bassin houil. d. Valen., -p. 368, pi. lviii. figs. 1-7. 

1911. ,, ,, Jongmans, Anleitung, i. p. 204, figs. 163-168. 

1869. Galamocladus equisetiformis, Schimper, Traite d. paleont. vcget., vol. i. p. 324, pi. xxii. figs. 1, 2, 3. 

1836. Hippurites longifolia, L. ifc H., Fossil Flora, vol. iii. pis. cxc, cxci. 

1876. Calamostachys, Boulay, Terr, houil. du Nord de la France et ses vegetaux foss., p. 24, pi. i. 

figs. 2, 2 bis. 
1876. Calamostachys germanica, Weiss, Steink. Galamar., part i. p. 47, pi. xvi. figs. 3, 4. 

Note. — -Very plentiful. Both sterile branchlets and cones. 
Horizons and Localities. — 

Blue Measures, above Brooch Coal: Jubilee Pit, Sand well Park, West 

Bromwich ; Hamstead Colliery, Great Barr, near Birmingham. 
Roof of Brooch Coal : Himley ; Shut End ; Pensnett ; Bilston ; Holly Hall, 

near Dudley. 
Brooch Binds Ironstone : Pensnett. 
Roof of Thick Coal: Bradley Colliery, Bilston. 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley ; 

Ettingshall. 
Heathen Coal : Dudley. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton ; 

Ruiton, near Sedgley. 

Asterophyllites longifolius Sternb., sp. 

1826. Bruckmannia longifolia, Sternb., Essai fiore monde prim., vol. i. fasc. iv. p. xxix and p. 50, pi. 

lviii. fig. 1. 

1828. Asterophyllites longifolia, Brongt., Prodrome, p. 159. 
1831. „ ,, L. & H., Fossil Flora, vol. i., pi. xviii. 

1888. Asterophyllites longifolius, Zeiller, Flore foss. bassin houil. d. Valen., p. 374, pi. lix. fig. 3. 
1911. ,, ,, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 118. 

1911. „ „ Jongmans, Anleitung, vol. i. p. 214, figs. 175-177. 

Note. — Very rare. 

Horizon and Locality. — Ten -foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Asterophyllites grandis Sternb., sp. 

1826. Bechera grandis, Sternb., Essai flore monde prim., vol. i. fasc. iv. pp. xxx and 46, pi. xlix. fig. 1. 
1836. ,, ,, L. & H., Fossil Flora, vol. iii., pi. clxxiii. (non vol. i., pi. xix. fig. 1). 

1886. Asterophyllites grandis, Zeiller, Flore foss. bassin houil. d. Valen., p. 376, pi. lix. figs. 4-7. 
1911. ,, ,, Jongmans, Anleitung, vol. i. p. 224, tig. 185. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 121 

Note. — Rare. 

Horizons and Localities. — 

Brooch Binds Ironstone : Pensnett. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 

Asterophyllites charseformis Sternb., sp. 

1826. Bechera charxformis, Sternb., Essai flore monde prim., vol. i. fasc. iv. p. xxx, pi. lv. fig. 3 

fi g- 5 )- 

1845. Asterophyllites charxformis, Unger., Syn. plant, foss., p. 33. 

1911. ,, ., Kidston, Mem. Musee roy. d'hist. nat. de Belgique, p. 119, pi. xi. 

figs. 2, 3, 3a, 4, 5. 
1911. .. ., Jongmans, Anleitung, vol. i. p. 232, figs. 190, 191. 

Note. — Not very common. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Annularia Sternberg. 
Annularia radiata Brongt. 

1822. Asterophyllites radiatus, Brongt., Class, d. veget. foss., p. 35, pi. ii. figs, la and lb. 

1826. Annularia radiata, Sternb., Essai flore monde prim., vol. i. fasc. iv. p. 31. 

1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 394, pi. lix. fig. 8, pi. Ixi. 

figs. 1, 2. 
1911. „ „ Jongmans, Anleitung, vol. i. p. 252, figs. 206-208 (? fig. 209). 

Note. — Common. Under this name are included the foliage branches of Calamites 
ramosus Artis, but most probably also those of other species. 
Horizons and Localities. — 

Roof of Brooch Coal : Shut End, near Kingswinford ; Holly Hall, near Dudley, 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley ; 

Ettingshall. 
Roof of Thick Coed : Bradley Colliery, Bilston. 
. Roof of Fireclay Coal: Doulton's Clay Pit, Netherton, near Dudley. 

Between Fireclay Coal and Bottom Coal: Doulton's Clay Pit, Netherton, 
near Dudley. 

Annularia galioides L. & H., sp. 

1804. Parkinson, Organic Remains, vol. i. pi. v. fig. 1. 

1832. Asterophyllites galioides, L. & H., Fossil Flora, vol. i. p. 79, pi. xxv. fig. 2. 

1893. Annularia galioides, Kidston, Trans. Roy. Soc. Edin., vol. xxxvii. p. 317, pi. ii. fig. 4. 

1899. „ „ Zeiller, Etude sur la Hore foss. d. bassin houil. d'Heraclee, p. 63, pi. v. 

figs. 16, 17. 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 16 



122 DR ROBERT KIDSTON ON THE 

1886. Annularia mierophylla, Zeiller (non Sauveur), Flore foss, d. bassin houil. de Valen., p. 392, 

pi. lx. figs. 3, 4. 

1887. ,, „ Stur (non Sauveur), Galamaraen d. Carbon-Flora d.Schatzlarer Schichtert, 

p. 211, pi. xiv. figs. 8, 9; pi. xv.6, fig. 2. 
1907. „ ,, Zalessky (non Sauveur), Bull. Comite geol. (St Petersbourg), vol. xxvi. 

p. 429, pi. xviii. fig. 3. 
1869. Annularia minuta, Wood, Trans. Amer. Phil. Soc, vol. xiii. p. 347, pi. viii. fig. 2. 
1884. ,, ,, Lesqx., Goal Flora, vol. iii. p. 725, pi. xcii. fig. 8. 

Remarks. — Until quite recently I believed that Annularia mierophylla Sauveur # 
was synonymous with the Annularia galioides L. & H., sp. In this view I think 
I have been mistaken, and now identify as Sauveur's plant specimens that have been in 
my collection for some years, and which have been derived from different localities, but 
which I had hitherto failed to identify. 

It may be stated here that in Annularia mierophylla Sauveur the leaves are 
sickle-shaped and much narrower in proportion to their length than those of Annularia 
galioides L. & H., sp. 

Annularia mierophylla Sauveur does not occur in this coal field, so a description 
of that plant is given in an Appendix to this paper, along with figures for comparison 
with Annularia galioides L. & H. 

All the plants which in the past I have recorded under the name of Annularia 
galioides do, however, belong to that species, which is not common in the South 
Staffordshire Coal Field. 

Horizons and Localities. — 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Roof of Thick Coal : Bradley Colliery, Bilston. 

Annularia sphenophylloides Zenker, sp. 

1833. Galium sphenophylloides, Zenker, Neues Jahrb., p. 398, pi. v. figs. 6-9. 
1837. Annularia sphenophylloides, Gutbier, Isis, p. 436. 

1882. ,, ,, SterzeL Zeitsch. d. deut. geol. Gesell., vol. xxxiv. p. 685, pi. xxviii. 

figs. 1-10. 

1886. „ „ Zeiller, Flore foss. bassin houil. d. Valen., p. 388, pi. lx. figs. 5, 6. 
1911. ,, ,, Jongmans, Anleitung, vol. i. p. 260, figs. 211, 212. 

1887. Annularia sarepontana, Stur, Calamarien d. Carbon-Flora d. Sehatzlarer Schichten, p. 221, 

pi. xiii. b, fig. 1 j pi. xiii. 6 (bis), fig. 1. 

Cones. 

1876. Stachannularia ealathifera, Weiss, Steinlc. Calamarien, Heft i. p. 27, pi. iii. fig. 11. 
1884. Calamostachys, cf. ealathifera, Weiss, ibid., Heft ii. p. 178. 

Note. — Very rare. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

* Ve'ye't.foss. d. terr. houil. d. Belgique, pi. lxix. fig. 6, 1848. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 123 

Annularia stellata Schl., sp. 

PL IX. figs. 1 and la. 

1804. Schlotheim, Flora d. Vorwelt, p. 32, pi. i. fig. 4. 

1820. Casuarinites stellatus, Schloth., Petrefactenkunde, p. 397. 

1860. Annularia stellata, Wood, Proc. Acad. Nat. Sciences Phil., p. 236. 

1886. ,, „ Zeiller, Flore foss. bassin houil. d. Valen., p. 398, pi. lxi. figs. 3-6. 

1911. „ „ Jongmans, Anleitung, vol. i. p. 238, figs. 193-196, 200-203. 

1845. Annularia longifolia, Gerniar, Vers. v. Wettin u. Lbbejun, p. 25, pi. ix. figs. 1-4. 

Cones. 
1826. Bruckmannia tuberculata, Sternb., Essai flore monde prim., vol. i. fasc. iv. pp. 45 and xxix, 

pi. xlv. fig. 2. 
1876. Stachannularia tuberculata, Weiss, Steinkohlen Calamarien, Heft i. p. 17, pi. i. figs. 2-5, pi. ii. 

figs. 1-3 and 5 (left-hand figure), pi. iii. figs. 3-10, 12. 
1884. Calamostachys (Stachannularia) tuberculata, Weiss, ibid., Heft ii. p. 178. 

Remarks. — The largest whorls of leaves on this specimen are only 1*30 cm. in 
diameter, but notwithstanding their comparatively small size I have no hesitation 
in referring the fossil to Annularia stellata. The specimen is shown natural size 
on PL IX. fig. 1, and a portion enlarged 2 times at fig. la. The leaves are lanceolate 
spathulate and single-veined. Their widest part is about f of their length from the 
base, and they terminate in subacute or blunt points. They agree in form perfectly 
with the leaves of Annularia stellata Schl., sp., and only differ in size. This, however, 
may be explained by the specimen having occupied a position near the termination 
of a branch, or even possibly having been part of a young individual. 

In size the whorls are less than in Annularia sphenophylloides, var. intermedia, 
Lesqx., as figured by Sellards,* but in this plant the leaves are broadly spathulate 
with blunt apices, quite typical of Annularia sphenophylloides, whereas the leaves of the 
Sandwell specimen are typical in form with those of Annularia stellata. 

This and another specimen from the same locality are the only examples I have 
yet seen from the Westphalian of Britain, but it occurs very near the top of the series. 
Annularia stellata has, however, been recorded from the Westphalian Series of the 
Continent. 

Horizon and Locality. — Blue Measures, above Brooch Coal : Jubilee Pit, Sandwell 
Park, West Bromwich. 

Calamostachys Schimper. 

Calamostachys Solmsi Weiss. 

PL IX. figs. 4, 4a, 46, 4c, id. 

1876. Macrostachya infundibidiformis, var. Solmsi, Weiss, Steink. Calam., Heft i. p. 73, pi. xviii. fig. 1 

(pars), figs. 3, 3a, and 4. 
1884. Calamostachys Solmsi, Weiss, Steink. Calam., Heft ii. p. 177. 
1911. ,, ,, Jongmans, Anleitung, vol. i. p. 288, fig. 236. 

* Foss. Plants, Upper Paleozoic, Kansas, p. 425, pi. liii. fig. 5, 1908. 



124 Dit ROBERT KIDSTON ON THE 

Description.— Cone heterosporous, pedicellate, complete length and apical portion 
unknown ; cylindrical, of almost equal width, but gradually narrowing upwards and 
from 12 to 16 mm. wide, internodes from 5 to 7 "5 mm. long. Bracts about length of 
internode or slightly longer, broadly lanceolate, with a base about 1*3 mm. wide, outer 
surface longitudinally striated and slightly keeled. Sporangiophores placed on the 
axis mid-way between the bract verticils as in Calamostachys, but the form of sporangio- 
phore and attachment of sporangia unknown. Microsporangia and megasporangia 
occur on same verticil and throughout all regions of the cone. Megaspores n 573 to 
fj- 648 in diameter, with small triradiate ridge, outer surface smooth. Microspores 
/«115tOMl30in diameter, smooth, with small triradiate ridge. 

Remarks. — The cone is incomplete at the apex, but the portion preserved is 1 1 '50 cm. 
long, and is given natural size on PI. IX. fig. 4, and a portion enlarged 2 times at 
fig. 4a. It contracts at the base, where it is attached to a finely striated and, as far as 
preserved, unjointed pedicel 2 cm. long and 5 mm. wide. The broadest part of the 
cone is about 2 cm. above its base, where it is 1*60 cm. wide. From this point it 
gradually narrows upwards, and about 7 cm. from the base it is T40 cm. wide. Above 
this no satisfactory measurements can be made, as one of the margins is slightly covered 
by the matrix. 

The cone shows 19 internodes, which vary from 6 mm. in length at the base to 
about 5 mm. at the top of the specimen. 

The bracts are broadly lanceolate, about 7 mm. long, and terminate in sharp fine 
points which extend very slightly beyond the node next above them. They have a 
striated outer surface, and some of them show indications of a central keel, but no mid- 
rib is observable. 

There appear to have been 16 or 18 bracts in a verticil, as 8 or 9 are seen on 
the exposed surface of the cone. 

The sporangiophores are not seen on the specimen. 

Small portions of the carbonaceous remains of the cone were removed from the base, 
the middle, and from near the top of the specimen, which were treated by the 
maceration process, when in all cases, even though the material operated on was small, 
the samples yielded microspores and megaspores. It would appear, then, that each 
verticil bore both microsporangia and megasporangia, and that the two forms of spores 
were not restricted to a special region of the cone. 

Some microspores enlarged 50 times are given at PL IX. fig. 46, and one 
enlarged 500 times is seen at fig. 4c, and a megaspore enlarged 50 times is 
shown at fig. id. Both microspores and megaspores do not seem to have been fully 
ripe, as their walls are thin and much crumpled. Their outer surface is smooth, and 
they are provided with a small triradiate ridge. This is shown on the microspore at 
fig. 4c. On only one case could I discover the triradiate ridge on a megaspore, and it 
was feebly developed ; this, as well as their crumpled condition, indicates a somewhat 
early state of development. The occurrence in the same verticil, and at different 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 125 

heights on the cone, of megasporangia and microsporangia, as evidenced by the occur- 
rence of the two forms of spores, is an interesting character of this cone. 

This species, which was collected by Mr H. W. Hughes, has not previously been 
recorded in Britain. 

Horizon and Locality. — Roof of Neiv Mine Coal: Clattershall Colliery, Brettell 
Lane. 

Calamostachys sp. 

Among some specimens found at Netherton were fragments of a small Calamostachys. 
None of the cones were complete, they are only 2 5 mm. broad, including the 
spread of the arcuate bracts, and the internodes are about 1 mm. long. The bracts 
bend gently upwards, and are rather more than one internode long. It is larger than 
the Calamostachys ? nana Weiss, # and does not appear to belong to any known 
species ; but the material is too fragmentary for a satisfactory description. 

The specimens were collected by Mr H. W. Hughes. 

Horizon and Locality. — Between Fireclay Coal and Bottom Coal: Doulton's 
Clay Pit, Netherton, near Dudley. (K. No. 4620.) 

Calamostachys sp. 

Horizon and Locality. — Ten -foot Lronstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. (K. No. 824.) 

Palseostachya Weiss. 

Palseostachya Ettingshauseni Kidston. 

PL IX. figs. 3, 3o-3c. 

1854. Calamites cummunis, Ett. (pars), Steinlwhlenf. v. Radnitz, p. 24, pi. viii. figs. 1 and 4. 

1869. Volkmannia elongata, Roehl (non Presl), Foss. Flora Steink.-Form. Westph., p. 19, pi. vii. fig. 1. 

1869. Calamostachys typiea, Schimper (pars), Trait'e d. paleont. veget., vol. i. p. 328 (pi. xxiii. fig. i.?), 

vol. iii. p. 457. 
1890. ,, ,, Kidston, Trans. York. Nat. Union, pp. 14, 23. 

1909. Calamostachys ? typiea, Arber, Fossil Plants, p. 74, fig. 57. 
1884. Calamostachys Ludwiyi, Weiss (pars), Steinkohlen Calamarien, part ii. p. 163, pi. xviii. fig. 2 

(non pi. xxii. figs. 1-8, pi. xxiii., pi. xxiv.). 
1903. Paleeostachya Ettingshauseni, Kidston, Trans. Roy. Soc. Edin., vol. xl. part iv. p. 794. 
1911. „ ,, Jongmans, Anleitung, vol. i. p. 327, figs. 284-286. 

1887. Calamites Sachsei, Stur (pars), Calamarien d. Schatz. Schichten, pp. 188, 189, pi. ii. fig. 7, 

pi. xi. fig. 7. 

Description. — Cone cylindrical, short-stalked, verticillate, apparently only four 
in each whorl, the panicle ending in a terminal cone ; cones attaining a length of 
7 cm. with a width of 80 mm. Internodes short, about 3. mm. long; bracts arcuate, 

* Steinkohlen Calamarien, Heft ii. p. 175, pi. xxi. fig. 10. 



126 DR ROBERT KIDSTON ON THE 

slightly spreading, apex sometimes incurved, and when perfect about the length of 
two internodes, more numerous than the sporangiophores. Probably six sporangio- 
phores in a whorl, which arise from the axil of a bract, each bearing four sporangia ; 
sporangia oval, homosporous. Microspores m 70 to m 100 in diameter, smooth, with a 
small triradiate ridge. 

Remarks. — The cone shown on PL IX. fig. 3, natural size, wants a small part 
of both the apex and the base, but is otherwise very well preserved. 

It is slightly contracted at the base, and probably little is missing from that end. 
The part preserved is 620 cm. long and about 0*80 cm. wide measured across the 
sporangia, but the bracts extend further outwards. It contains eighteen short 
internodes about 3*75 mm. long. Unfortunately, the specimen does not enable one 
to determine the number of the lanceolate bracts which compose the whorls. They 
at first spring outwards at almost right angles to the axis, and then bend upwards 
past the spreading sporangia which spring from their axils and reach almost to the 
top of the second internode above that from which they spring. 

The sporangia are large, oval, slightly narrowed at base, and from 270 mm. to 
3 '10 mm. long, and therefore occupy almost the whole length of the internode. It is 
difficult to determine the number of sporangial groups in a whorl, owing to the 
compressed condition of the specimen, but I do not think the sporangiophores were 
very numerous, perhaps not more than six in a whorl. If one examines the portion 
of the cone enlarged 2 times at fig. 3a, there do not appear to be more than three 
pairs of sporangia visible on the exposed surface of the cone ; so we are therefore 
probably looking at the exposed surface of three groups of four sporangia. In the 
whole verticil there would therefore probably be six sporangiophores, each bearing 
four sporangia. 

One of the sporangia from the base of the cone was removed and treated 
by the acid-ammonia process, when it was found to contain very numerous circular 
microspores with a small triradiate ridge, measuring on an average /« 80 in diameter. 
Palseostachya Ettingshauseni is therefore probably homosporous. Some microspores 
are seen enlarged 50 times at fig. 36, and one 500 times at fig. 3c. 

Palseostachya Ettingshauseni, which is the cone of Catamites Sachsei Stur (pars), 
is closely related to Palseostachya elongata Presl, sp., but is easily distinguished 
by its small size and less dense mode of growth. 

It is not common in the South Staffordshire Coal Field, where, though the cone 
has been found, no specimens of the parent plant. Calamites Sachsei Stur, have been 
discovered, as far as 1 am aware. 

The specimen figured by Lindley and Hutton on pis. xv. and xvi. of their Fossil 
Flora as the foliage of Calamites nodoaus (non Schlotheim) ( = Calamites ramosus Artis) 
are fortunately preserved. The small branch shown on pi. xv. has no organic con- 
nection with the stem which occurs close to it, and its position is merely accidental. 
Both this and the specimen given on pi. xvi. are fertile branches of Palseostachya 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 127 

Ettingshauseni ; the latter figure only shows one of several cone-bearing branches 
which occur on the slab. 

The originals of these figures are preserved in the " Hutton Collection," Hancock 
Museum, Newcastle-on-Tyne. 

Horizon and Locality. — Roof of New Mine Coal: Mount Pleasant, Brierley Hill. 

Palseostachya elongata Presl, sp. 

1838. Volkmannia elongata, Presl, Verhandl. d. Gesell. d. vaterldndisehen Museums in Bbhmen, p. 27, 

pi. i. (fide Weiss). 
1872. ,, ,, Feistm., Abhandf. d.k. bohm. Gesell. d. Wissensch., vi. Folge, vol. v. p. 20, 

pi. iv. fig. 3, pi. v. fig. 2. 
1874. „ ,, Feistm., Vers. d. bohm. Ablager, i. Abth. p. 119, pi. xiii. figs. 1, 2. 

1876. Palxostacliya elongata; Weiss, Steink. Calamarien, part i. p. 108, pi. xv. ; part ii. p. 181, pi. xxii. 

fig. 15. 
1911. ,, . ,, Jongmans, Anleitung, vol. i. p. 324, figs. 281-2. 

Note. — Very rare. Collected by Mr H. W. Hughes. 
Horizons and Localities. — 

(?) Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
This specimen is fragmentary, and not in a suitable condition for a 
satisfactory determination. (K. No. 3700.) 
Roof of New Mine Coal : Merryhill Colliery, Mount Pleasant, Brierley Hill. 

Palseostachya gracillima Weiss. 

1884. Palxostachya gracillima, Weiss, Steinkohlen-Calamarien, part ii. p. 184, pi. xviii. fig. 1. 
1886. ,, „ Kidston, Trans. Geol. Soc. Glasgoio, p. 54, pi. iii. fig. 3. 

1910. ,, „ Arber, Proc. Yorkshire Geol. Soc, vol. xvii. part ii. p. 143, pi. xii. 

1911. ,, „ Jongmans, Anleitung, vol. i. p. 326, fig. 283. 

Remarks. — This species is rare in the South Staffordshire Coal Field, and I only 
know of the plant having been collected once by Mr H. W. Hughes, F.G.S. 

Horizon and Locality. — Roof of Fireclay Coal: Doulton's Clay Pit, Netherton. 

Palseostachya minuta Kidston, n. sp. 
PI. XI. figs. 1, la-lc. 

Description. — Cone small, 1*5 cm. long and about 2*5 mm. wide, with somewhat 
blunt apex, bracts arcuate, distal portion upright, apex slightly incurved, about 
1^ times as long as internodes. Internodes 1 to 1'25 mm. long. Probably six 
sporangiophores in a whorl. 

Remarks. — Some small slabs were collected by Mr H. W. Hughes, thickly covered 
with these cones, of which a slab is seen at fig. 1, PI. XL, natural size. They are 
detached from their parent stems and are usually more or less imperfect, though 



128 DR ROBERT KIDSTON ON THE 

a few are complete. On the exposed surface of the cones three groups of sporangia 
are frequently visible, and probably the fertile whorls contained six sporangiophores. 
In any case, their number must have been small. 

Some of the cones are enlarged 2 times at figs, la to \c to show their general 
character. 

Horizon and Locality. — Between Fireclay Coal and Bottom Coal: Doulton's 
Clay Pit, Netherton, near Dudley. (K. No. 4476.) 

Huttonia Sternberg. 

Huttonia spicata Sternb. 
PL XL fig. 4, PL XIV. fig. 4. 

1837. Huttonia spicata, Sternb., Verhandl. d. Gesellsch. d. vaterl. Museums, Bbhmen, 1837, p. 69, pi. i. 

figs. 1-4 {fide Jongmans). 
Andrae, in Germar, Vers. v. Wettin u. Lobejun, p. 91, pi. xxvii. fig. 4. 
Feistmantel, Abhandl. d. k. bohm. Gesell. d. Wissensch., vi. Folge, vol. v. p. 7, 

pi. i. fig. 1 (" Ueber Fruchtstadien fossiler Pflanzen aus der bb'hmischen 

Steinkohlenformation "). 
Feistmantel, Vers. d. bohm. Ablager, Heft i. p. 113, pi. viii. fig. 3. 
Weiss, Steinkohl. Calamarien, part i. p. 82, pi. xiii. figs. 3, 4 ; pi. xiv. figs. 1-4. 
Schenk, in Richthofen, China, p. 234, pi. xli. figs. 1-3. 
Weiss, Steinkohl. Calamarien, part ii. p. 188, pi. xxi. fig. 9. 
Jongmans, Anleitung, vol. i. p. 352, figs. 320-324. 

Description. — Cones with short unjointed stems, leafless, bracts with a small base, 
enlarging upwards and ending in a suddenly contracted long sharp point, the sides of 
which are convex, 18 to 20 in a whorl, with outer surface longitudinally striated. The 
bracts of the lowest whorls are smaller and more truly lanceolate. Bracts free to the 
base ; and situated below them is a pendulous sheath. The sporaDgiophores spring 
from the axils of the bracts and are directed outwards. 

Remarks. — The specimen is shown natural size at PL XI. fig. 4, and a portion 
enlarged 2 times at PL XIV. fig. 4. This latter figure shows well at the broken- 
over upper end the narrowed basal portion of the bracts, whose upper surface is 
longitudinally striated. The fossil is only a small part of the cone, of which I am not 
aware that a complete example has ever been discovered. It is extremely rare to find 
the bracts complete, the upper portion being almost always broken off, as in our 
solitary example from South Staffordshire. The internodes are short, about 4'50 mm. 
long, and the bracts, which appear to have been almost coreaceous, show no indication 
of a midrib. This, which is the first record of Huttonia spicata for Britain that has 
come under my notice, was collected by Mr H. W. Hughes, F.G.S., to whom I am 
so much indebted for my knowledge of the fossil plants of the South Staffordshire 
Coal Field. 

Horizon and Locality. — Blue Measures, six feet above Fireclay Coal : Doulton's 
Clay Pit, Netherton, near Dudley. 



1851. 


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1875. 


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FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 129 

Sphenophyllese. 

Sphenophyllum Brongniart. 
Sphenophyllum cuneifolium Sternb., sp. 

1823. Rotularia cuneifolia, Sternb., Essai flore monde prim., vol. i. faso. ii. ppi 33 and 37, pi. xxvi. 

figs. 4a, 45. 
1886. Sphenophyllum cuneifolium, Zeiller, Flore foss. bassin houil. d. Valen., p. 413, pi. lxii. fig. 1, 

pi. lxiii. figs. 1-10. 
1893. ,, ,, Zeiller, Mem. Soc. geol. d. France: Paleont., vol. iv. No. ii. p. 12, 

pi. i. (iii.) figs. 1-4, pi. ii. (iv.) figs. 1-3, pi. iii. (v.) figs. 1, 2. 
1901. ,, ,, Kidston, Trans. Nat. Hist. Soc. Glasgow, vol. vi. (new series) p. 124, 

fig. 21, a, b, p. 121, and fig. 23, p. 125. 
1911. ,, ,, Jongmans, Anleitung, p. 377, figs. 335-345a (non 345 b, c) (non 

" nach Kidston "). 
1826. Rotularia saxifragsefolium, Sternb., Essai flore monde prim., vol. i. fasc. iv. p. xxxii, pi. Iv. fig. 4. 
1831. Sphenophyllum erosum, L. & H., Foss Flora, vol. i., pi. xiii. 

Note. — Common. 
Horizons and Localities. — 

Brooch Coal : Holly Hall, near Dudley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley ; 

Cabbage Hall Pit, Netherton, near Dudley. 
Roof of New Mine Coal: Brierley Hill; Dibdale, near Gornal. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton, near 
Dudley. 

forma saxifragsefolium Sternb., pro. sp. 
Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr, 

near Birmingham. 
Brooch Coal : Oldbury. 

Roof of New Mine Coal: Merry hill Colliery, Brierley Hill. 
Blue Measures, six feet above Fireclay Coal: Netherton, near Dudley. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton, near 

Dudley. 

Sphenophyllum tenuissimum Kidston, n. sp. 
PI. XVI. figs. 3, 4, and 4a, 5. 

Description. — Stem very slender, about 1 mm. wide, internodes 7 mm. long. 
Leaves single veined, 8 mm. long or more, narrow, about 1 mm. broad, dichotomously 
divided, and segments end in an acute point. Fructification borne on the ordinary 
stems without any cone formation, and consisting of numerous pyriform sporangia, 
united to each other by a thick basal extension into groups of four. 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 17 



130 



DR ROBERT KIDSTON ON 'THE 



Remarks. — This extremely delicate species belongs to the " Selago" type, in which 
the sporangia are borne on little altered or unmodified portions of the stem, as in 
Sphenophyllum majus Bronn, sp., # and Sphenophyllum charaeforme Jongmans.f 

The sporangia are pyriform, and, though they may have combined in other numbers, 
those seen here, where their number can be determined, are united in groups of four. 
The individual sporangia are very small, including the basal stalk-like portion by which 
they are united to each other ; they are only 1 mm. long, and slightly narrower. The 
groups are much broken up ; but at the uppermost node, seen in text-fig. 6, which is 




Text-fig. 6. — Sphenophyllum tenuissimum Kidston. Specimen enlarged 4 times. 



enlarged 4 times, two perfect and one imperfect sporangium are seen united to each 
other, and the point from which the fourth has been removed is also seen. A photograph 
of this group, enlarged about 3 times, is given on PI. XVI. fig. 5, where the granular 
appearance of the sporangia represents the roughness of the matrix and is not 
organic. 

The leaves are long for the size of the stem. None are perfect, but at text-fig. 6, 
&', a bifurcation of the limb is seen, and the one remaining perfect segment ends in a 
sharp point. At b" another fragment of a leaf is preserved, which has also dichotomised. 
Although one cannot speak with certainty, the leaves probably only forked once, and 
bore the sporangia on their upper surface in the neighbourhood of the stem, as in 

* Kidston, Mem. Muse'e roy. d'hist. nat. de Belgique, vol. iv. p. 221, pi. xiv. figs. 1-4, 4a ; pi. xv. figs. 2, 3, 1911. 
t AanahiL d. L. It. /taturhist. Ho/museums. Wien, vol. xxvi. p. 449, pi. vi. and four text-figs., 1912. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 131 

Sphenophyllum majus, the only other known species in which the sporangia were united 
in groups of 4 or 6. 

In Sphenophyllum charxforme Jongmans, another very delicate species, the 
sporangia appear to be united singly to a sporangiophore, as in Sphenophyllum 
cuneifolium Sternb., sp., and this at once separates Sphenophyllum tenuissimum from 
that species. 

The specimen, which is preserved in an ironstone nodule, is seen natural size on 
PL XVI. figs. 3, 4. It was collected by Mr H. W. Hughes. 

Horizon and Locality.' — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Sphenophyllum sp. 
PI. X. figs. 5, 5a. 

1911. " Microsporanges," Carpentier, " Sur quelques fructifications et inflorescences du "Westphalien du 
Nord de la France," Revue generate de bot., vol. xxiii. p. 11, pi. xiv. figs. 6, 7. 

Remarks. — Similar groups of sporangia to those figured by M. l'Abbe Carpentier 
from the Nord Coal Field, France, occur in the Yorkshire and South Staffordshire Coal 
Fields ; the former were collected by Mr W. Hemingway, and the latter by Mr H. W. 
Hughes. They consist of four or five obtusely pyriform, or what might be almost 
better described as " fig-shaped," sporangia, united in groups of four or five by their 
contracted pedicel-like bases. A quadrate group of these sporangia is shown at fig. 5, 
PI. X., natural size, and enlarged 2 times at fig. 5a. The diameter of the group is 
about 4 mm. 

The sporangial termination is as broad as or even slightly broader than long, and its 
apex sometimes shows a faint indication of a notch. 

Abbe Carpentier compares these sporangia to the fructification of Telangium 
(Sorocladus) stellatum Lesqx., sp.,* but with this type of fructification I do not think 
they have any relationship. 

The only plants with which I am acquainted that have the same type and arrange- 
ment of sporangia are Sphenophyllum majus Bronn, sp.,f and Sphenopyhyllum 
tenuissimum, described here. 

From the sporangia of the latter they are at once distinguished by their much larger 
size, and from the former they differ in being flattened at their apex and broader in 
proportion to their length, and have more the form of a fig than of a pear ; but Dr Paul 
Bertrand, J with whom I have had some correspondence on these small fossils, mentions 
that his specimens, of which he has shown me excellent photographs, and which are 
identical with that figured here, have discharged their spores, and he suggests that 

* Lesquereux, Coal Flora, p. 328, pi. xlviii. fig. 9, 1879. Zeiller, Flore foss. bassin houil. d. Valen., p. 141, pi. xii. 
fig. 2, 1886. 

t See Kidston, Mem. Muse'e roy. d'hist. nat. de Belgique, vol. iv. p. 221, pi. xiv. figs. 1-4, 4a ; pi. xv. figs. 2, 3 ; and 
text-fig. 35. 

+ In letter of October 19, 1913. 



132 DR ROBERT KIDSTON ON THE 

this may be the cause of their difference in shape from those on the fertile specimens 
of Sphenophyllum majus. I scarcely think that this explanation will explain their 
difference in form, but nevertheless they are very probably from a plant closely 
related to Sphenophyllum majus, but I do not think we can refer them to that species, 
though 1 know of no Sphenophyllum from the same horizon to which they might belong. 
Horizon and Locality. — Blue Measures, above Brooch Coal : Jubilee Pit, Sandwell 
Park, West Bromwich. 

Lycopodiacese. 

Lycopodites Brongniart (pars). 

Lycopodites Meeki Lesqx. 

1870. Lycopodites Meekii, Lesqx., Geol. Survey of Illin., vol. iv. p. 426, pi. xxvi, figs. 6 and 6a. 
1874. ,, „ Schimper, Traite d. paleont. veget., vol. iii. p. 532. 

1879. ,, ,, Lesqx., Coal Flora, p. 857, pi. lxii. figs. 1 and la. 

Remarks. — Mr Henry Insley has collected a slab, about 21 cm. broad by 25 cm. 
long, bearing many branchlets of this delicate Lycopod. The longest branch, which 
bifurcates four times at an acute angle, has a length of about 2 1 cm. , and at its broken- 
over lower extremity it measures about 5 mm. across. The surface of the stem bears 
fusiform cushions very similar in form to those of Lepidodendron, about 5 mm. long 
and 0'75 mm. wide. The leaves are very narrow, almost setaceous, and their greatest 
length is about 175 cm. They are occasionally adpressed, but more commonly they 
spread slightly outwards from the stem at an acute angle. 

A midrib has not been observed in the leaves. 

As Lesquereux has already pointed out, Lycopodites Meeki has a general 
resemblance to the Lepidodendron selaginoides Sternb., especially to his pi. xvii. 
fig. 1, # but is much more slender and the leaves narrower and longer in proportion to 
their width. 

In the absence of fructification there may be some doubt as to the systematic 
position of this plant, but since many of the species formerly included in Lycopodites 
are now placed in Selaginellites Zeiller,t it leaves Lycopodites Brongniart (pars) free 
for the retention of such imperfectly known species as the Lycopodites Meeki Lesqx. 

Horizon and Locality. — Blue Measures, six feet above Brooch Coal: Hamstead 
Colliery, Great Barr, near Birmingham. 

Lepidodendron Sternberg. 

Lepidodendron ophiurus Brongt. 

PL XL figs. 2, 3. 

1822. Sagenaria ophiurus, Brongt., Class, d. veget. foss., pp. 27 and 90, pi. iv. figs, la and 16. 

1828. Lepidodendron ophiurus, Brongt., Prodome, p. 128. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 458, pi. lxviii. figs. 1-6. 

* Sternb., Essaiflore monde prim., vol. i. fasc. ii. pp. 29 and 35 ; fasc. iv. p. viii, pi. xvi. fig. 3, pi. xvii. fig. 1. 
+ Bassin houil. et perm, de Blanzy et du Creusot, p. 140, 1906. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 133 

1911. Lepidodendron ophiurus, Kidston, Mem. Musee roy. d'hist. not. de Belyique, p. 140. 
1831. Lepidodendron graeile, L. & H., Foss. Flora, vol. i., pi. ix. 
1838. ,, „ Brongt., Hist. d. veget. foss., vol. ii., pi. xv. 

1831. Lepidodendron dilatatum, L. & H., Fossil Flora, vol. i., pi. vii. fig. 2. 

1894. Lepidodendron elegans, Nathorst (non Sternb.), Palaeoz. Flora d. arktischen Zone, p. 34, pi. xvi. 
fig. 1 (Kongl. Svenska Vetenskaps-Akad. Handl., vol. xxvi. No. 4). 

Remarks. — Two specimens of the cones of Lepidodendron ophiurus are given 
natural size at figs. 2, 3, PL XL That at fig. 2 shows a cone, 14 cm. long and 1*50 cm. 
broad, of almost equal width throughout, terminating a branchlet which is easily 
identified by the presence of the characteristic foliage of Lepidodendron ophiurus. 

The distal portion of the lanceolate bracts is upright and mostly adpressed to the 
cone, and in this it differs from Lepidostrobus squarrosus Kidston,* where the distal 
portions of the bracts are spreading and the cone is also larger. 

The other specimen, that at fig. 3, and which is 15 cm. long, though not attached 
to the branch, from its size, form, and character of the bracts I have no hesitation in 
referring to the same species. This specimen is preserved in the " round " and has 
been thoroughly impregnated with carbonate of lime, now bleached white, while the 
surrounding matrix of clay ironstone is of a red-brown colour. The vegetable matter 
is converted into black carbon, so that the general structure of the cone stands out 
with diagrammatic clearness. 

At the base of the cone, the sporangia! portion of the bract is slightly deflexed, 
about 2 cm. above the base it becomes horizontal, and above this point it gradually 
assumes an upward direction. A slight contraction has taken place in this specimen, 
as shown by the cavity at the apex, originally occupied by the plant, being now filled 
with lime, though the apex of the cone itself is quite complete. 

Towards the centre of the cone given at fig. 3 a small part of the peripheral region 
is seen, and on this are exhibited fragments of some of the bracts. Neither the 
sporangia nor spores are preserved in this example. 

On another similarly preserved specimen in my collection f it is shown that the 
bracts were spirally placed on the axis. 

That given at fig. 2 was received from Sir Charles Holcroft, and that at fig. 3 
from Mr H. W. Hughes. 

Horizons and Localities.. — 

Roof of Brooch Coal : Shut End ; Himley ; Pensnett ; Holly Hall, Dudley ; 

Coneygre Colliery, Tipton. 
Brooch Binds Lronstone : Pensnett. 
Ten-foot Lronstone Measures : Clayscroft Openwork, Coseley, Dudley ; Tipton ; 

Moxley. 
Shale over Heathen Coal : Dudley. 
Gubbin Lronstone Measures : Shut End. 
Roof of New Mine Coal: Merry hill Colliery, Brierley Hill. 

* Trans. Roy. Soc. Edin., vol. xxxvii. p. 342, pi. iv. figs. 13, 14, 1893. t No. 3736. 



134 DR ROBERT KIDSTON ON THE 

Lepidodendron simile Kidston. 

1911. Lepidodendron simile, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 137. 

1837. Lepidodendron elegans, Brongt. (non Sternb.), Hist. d. vegef. foss., vol. ii. p. 35, pi. xiv. 

1880. Lepidodendron lycopodioides, Zeiller (non Sternb.), Veget.foss. d. terr. houil. de la France, p. 3, 

pi. clxxi. 
1882. ,, „ Renault (non Sternb.), Cows d. bota?i. foss., vol. ii. p. 14, pi. v. 

% 8. 
1886. „ ,, Zeiller (non Sternb.), Flore foss. bassin houil. d. Valen., p. 464, 

pi. lxix. figs. 2, 3, pi. lxx. fig. 1. 

Note. — Rare. 

Horizons and Localities. — 

Roof of New Mine Coal: Merryhill Colliery, Brierley Hill. 
Shale overlying Pont Hill Trap (probably corresponding in position to shales 
lying between the Fireclay Coal and the Bottom Coal) : Bentley Quarry, 
Walsall. Collected by Mr S. Priest. 

Lepidodendron aculeatum Sternb. 

1820. Lepidodendron aculeatum, Sternb., Essai flore monde prim., vol. i. fasc. i. pp. 21 and 25, pi. vi. 

fig. 2, pi. viii. fig. 1b, a, b ; fasc. ii. p. 28, pi. xiv. figs. 1-4 ; fasc. iv. 

p. 10. 
1886. ,, ,, Zeiller, Veget.foss. bassin houil. d. Valen., p. 435, pi. lxv. figs. 1-7. 

1858. Lepidodendron modulatum, Lesqx., in Rogers, Geol. of Pennsyl., vol. ii. p. 874, pi. xv. fig. 3. 

Note. — Common. 
Horizons and Localities. — 

Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr, 

near Birmingham. 
Roof of Thick Coal : Bradley Colliery, Bilston ; Clayscroft Openwork, Coseley, 

near Dudley. 
Below Thick Coal : Near Dudley. 

Gubbin Ironstone : Himley ; Bilston ; Eagle Colliery, Old Hill ; Clayscroft 
Openwork, Coseley, near Dudley. 

forma modulatum Lesqx., pro. sp. 
Roof of New Mine Coal: Doulton's Clay Pit, Netherton, near Dudley. 

Lepidodendron obovatum Sternb. 

1820. Lepidodendron obovatum, Sternb., Essai flore monde prim., vol. i. fasc. i. pp. 21 and 25, pi. vi. 

fig. 1, pi. viii. fig. 1a, a, b ; fasc. iv. p. 10. 
1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 442, pi. Ixvi. figs. 1-8. 

1858. Lepidodendron clypeatum, Lesqx., in Rogers, Geol. of Pennsyl., p. 875, pl. xv. fig. 5, 
pi. xvi. fig. 7. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 135 

Note. — Common. 
Horizons and Localities. — 

Roof of Brooch Coal : Himley ; Shut End. 

Gubbin Ironstone : High Haden Colliery, Old Hill ; Shut End ; No. 1 1 Pit, 

Woodside Colliery, Hartshill ; Clayscroft Openwork, Coseley, near Dudley. 

Shale overlying Pont Hill Trap (probably corresponding to shales between 

Fireclay and Bottom Coals) : Bentley Quarries, Walsall. Collected by 

Mr S. Priest. 



Lepidodendron acutum Presl, sp. 

1838. Bergeria acuta, Presl, in Sternb., Vers., vol. ii. fasc. vii.-viii. p. 184, pi. xlviii. figs, la, \b. 
1911. Lepidodendron acutum, Kidston, Mem. Musee roy. d'hist. not. d. Belgique, vol. iv. p. 146. 
1854. Lepidodendron Haidingeri, Ettingshausen, Steinkf. v. Radnitz, p. 55, pis. xxii., xxiii. 
1886. ,, „ Zeiller, Flore foss. bassin houil. d. Valen., p. 461, pi. lxix. fig. 1. 

1875. Sagenaria elegans, Feistm. (non Sternb.), Vers. d. bbhm. Ablager., ii. Abth. p. 29, pi. viii. figs. 3, 3a. 
1899. Lepidodendron elegans, Hofmann and Ryba {non Sternb.) (pars), Leitpflanzen, p. 80, pi. xiv. 
figs. 2, 3 (non pi. xiv. fig. 1, pi. xv. fig. 9). 

Note. — Not common. 

Horizon and Locality. — Roof of Fireclay Coal: Doulton's Clay Pit, Netherton, 
near Dudley. 



Cf. Lepidodendron distans Lesqx. 
[Lepidodendron serpentigerum Konig, Icones foss. sectiles, pi. xvi. fig. 195.) 

1858. Lepidodendron distans, Lesqx., Geol. of Pennsyl., vol. ii. p. 874, pi. xvi. fig. 5. 

1879. „ „ Lesqx., Goal Flora, vol. ii. p. 387, pi. lxiv. fig. 10. 

1858. Lepidodendron oculatum, Lesqx., Geol. of Pennsyl., vol. ii. p. 874, pi. xvi. fig. 4. 

1869. Lepidodendron chilallaeum, Wood, Trans. Amer. Phil. Soc, vol. xiii. p. 346, pi. ix. fig. 4. 

Remarks. — The specimen figured by Mr B. Smith in the Geol. Mag., May 1905, 
p. 209, as Lepidodendron sp. is most probably an aged example of Lepidodendron 
distans Lesqx. ( = Lepidodendron serpentigerum Konig). The characters which he 
appears to regard as precluding the reference of his specimen to this species may, I 
think, be ascribed to age and imperfect preservation, which in this case makes a satis- 
factory determination of his fossil impossible. The locality given for the specimen is 
"from the Middle Coal Measures near Dudley." It is contained in the Johnson 
Collection, British Museum (No. V. 1233). 

This species in my previous lists was included under the name of Lepidodendron 
serpentigerum, Konig, but it has been shown that the part of Konig's work containing 
the figure and name of Lepidodendron serpentigerum, though privately distributed, 
was never issued to the public* Konig's name must therefore give place to that of 
Lesquereux. 

* Zeiller, Flore foss. bassin houil. d. Valen., p. 708 (footnote), 1888. 



136 DR ROBERT KIDSTON ON THE 

LepidostrobuS Brongniart. 

Lepidostrobus variabilis L. & H. 

1831. Lepidostrobus variabilis, L. & H., Fossil Flora, pi. x., pi. xi. (fig. on right only). 

Remarks. — Very common in the ironstone nodules. 

The Lepidostrobus ornatus, L. & H., Fossil Flora, vol. i., pi. xxvi., can only be 
regarded as a cone of the Lepidostrobus variabilis type preserved "in the round" in 
ironstone uodules, where the distal portions of the bracts have decayed. Such specimens 
are not uncommon at Coseley. The authors of the Fossil Flora evidently misinterpreted 
their specimen. 

Horizons and Localities. — 

Blue Measures, above Brooch Coal : Hamstead Colliery, Great Barr, near 

Birmingham. 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton, near 
Dudley. 

Lepidostrobus Geinitzi Schimper. 

1855. Lepidostrobus variabilis, Geinitz (non L. & H.), Vers. d. Steinkf. in Sachsen, p. 50, pi. ii. figs. 

1, 3, 4. 

1869. „ „ Roehl {non L. & H.) (pars), Foss. Flora d. Steink.-Form. Westph., 

p. 142, pi. vii. fig. 2. 
1875. ,, „ Feistmantel (non L. & H.) (pars), Vers. d. bohm. Ablager., part ii. p. 44, 

pi. xiv., pi. xv. figs. 1, 2. 

1870. Lepidostrobus Geinitzi, Schimper, Traite de paleont. veget., vol. ii. p. 62, pi. lxi. fig. 6 (fig. 7?). 

Excl. syn. L. comosus L. & H. 
1886. ,, „ Zeiller, Flore foss. bassin houil. de Valen., p. 501, pi. lxxvi. fig. 2. 

1890. „ „ Renault, Flore foss. bassin houil. de Gommentry, part ii. p. 527, pi. lxi. 

figs. 5, 6. 

Horizon and Locality. — Blue Measures, six feet above Brooch Coal : Hamstead 
Colliery, Great Barr, near Birmingham. Collected by Mr H. Insley. 



Lepidostrobus triangularis Zeiller, sp. 

1911. Lepidostrobus triangularis, Kidston, Mem. Musee roy. d'hist. not. de Belgique, vol. iv. p. 158. 
1886. Lepidophyllum triangulare, Zeiller, Flore foss. bassin houil. d. Valen., p. 508, pi. lxxvii. 

figs. 4, 5. 
1897. Lepidophyllum Pichleri, Kerner, Jahrb. d. k. k. Reiehsanst., Band xlvii. Heft ii. p. 383, pi. x. 

figs. 1, 2 ("Carbonflora d. Steinacherjoches "). 

Remarks. — This species is closely related to Lepidostrobus anthemis Konig, sp., # 
the Lepidostrobus radians of Schimper,! and, if really distinct from it, differs only in 
the sides of the bracts being perhaps somewhat more concave. 

* Icones fossilium sectiles, pi. xvi. tig. 200. t Traite d. paldont. vfyet., vol. ii. p. 63. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 137 

Horizons and Localities. — 

Above Brooch Coal : Holly Hall, Dudley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley. 

Lepidophyllum Brongniart. 
Lepidophyllum intermedium L. & H. 

1832. Lepidophyllum intermedium, L. & H., Fossil Flora, vol. i. pi. xliii. fig. 3. 

1877 Lepidophyllum lanceolatum, Lebour (non L. & H.), lllustr. of Fossil Plants, p. 105, pi. liii. 

1890. Lepidophyllum magus, Renault (non Brongt.), Flore foss. terr. houil. de Commentry, part ii. 

p. 516, pi. lix. figs. 8, 9. 
1904. ,, ,, Zalessky (pars), Mem. d. Oomite geol. (St Petersbourg), nouv. ser., 

livr. xiii. p. 104, pi. vii. fig. 6 ( ? fig. 9). 

Note. — Rare. 

Horizons and Localities. — 

Roof of Brooch Coal: Tividale. 

Roof of Bottom Coal: No. 120 Pit, Coneygre Colliery, Tipton. 



LepidophloiOS Sternberg. 

Lepidophloios laricinus Sternb. 

1820. Lepidodendron laricinum, Sternb., Essai flore monde prim., vol. i. fasc. i. pp. 23 and 25, pi. xi. 

figs. 2-4. 
1826. Lepidophloyos laricinum, Sternb., ibid., vol. i. fasc. iv. p. 13. 
1855. Lepidophloios laricinum, Goldenberg (jiars), Flora Sarxp. foss., Lief. i. p. 22, pi. iii. figs. 14, 14a; 

Lief. iii. p. 30, pi. xv. fig. 11 (1 non fig. 9), pi. xvi. figs. 2, 3 (? 1, 4, 5, 6) (non figs. 7, 8), 

1862. 
1886. Lepidophloios laricinus, Zeiller, Flore foss. hassin houil. d. Valen., p. 471, pi. lxxii. figs. 1-3. 
1893. „ „ Kidston, Trans. Roy. Soc. Edin., vol. xxxvii. p. 155, pi. i. figs. 4, 4a ; 

pi. ii. figs. 8, 8a, 8b. 
1866. Lepidophloios Acadianus, Dawson, Quart. Journ. Geol. Soc, vol. xxii. pp. 163, 168, pi. x. 

figs. 45 a-h, figs. 50, 50a to c (non pi. xi. fig. 51). 

Note. — Rare. 

Horizon and Locality. — Roof of New Mine Coal : Clattershall Colliery, Brettell 
Lane. 

Halonia L. & H. 

Halonia tortuosa L. & H. 

1833. Halonia (?) tortuosa, L. & H., Fossil Flora, vol. ii. pi. lxxxv. 

1837. Halonia regularis, L. & H., ibid., vol. iii. pi. ccxxviii. 

1838. Halonia tuberculata, Brongt., Hist. d. veget. foss., vol. ii. pi. xxviii. figs. 1-3. 

Remarks. — Under this species are placed decorticated fertile branches of Lepido- 
phloios which cannot be referred to their respective species. 

TRANS. ROY. SOC. EDIN., VOL. L., PART I. (NO. 5). 18 



138 DR ROBERT KIDSTON ON THE 

Horizons and Localities. — 

Shale over Brooch Coal: Holly Hall, Dudley. 

White Ironstone : Shut End. 

Measures near roof of Bottom Coal: Powk Hill Quarry, Bentley, near Walsall. 

Bothrodendron L. & H. 
Bothrodendron minutifolium Boulay, sp. 

1876. Rhytidodendron minutifolium, Boulay, Terr, liouil. du Nord de la France, p. 39, pi. iii. figs. 1 and 

1 bis. 
1886. Bothrodendron minutifolium, Zeiller, Flore foss. bassin liouil. d. Valen., p. 491, pi. lxxiv. figs. 2-4. 
1889. ,, „ Kidston, Trans. Roy. Soc. Edin., vol. xxxv. p. 412, pi. ii. fig. 6. 

1889. ,, ,. Kidston, Proc. Roy. Phys. Soc. Edin., vol. x. p. 92, pi. iv. figs. 5, 6. 

1893. Siyillaria (Bothrodendron) minutifolium, Weiss, Die Siyillarien d. preuss. Steink. u. Rothl. 

Gebiete: II. Gruppe d. Subsigillarien, p. 49, pi. i. figs. 3, 4. ; pi. ii. figs. 8, 9. 
1893. Siyillaria minutifolium, var. rotundata, Weiss, ibid., p. 53, pi. i. fig. 6, pi. ii. fig. 7. 
1893. ,, ,, var. attenuata, Weiss, ibid., p. 53, pi. ii. figs. 10, 11. 

1831. Lepidodendron selaginoides, L. & H. (non Sternberg), Fossil Flora, vol. i. pi. xii. (plate very 

incorrect). 
1875. Lycopodium carbonaceum, Feistmantel, Vers. d. bbhm. Kohlenab., Abth. ii. p. 9, pi. i. figs. 1, 2. 
1886. Lepidostrobus olryi, Zeiller, Flore foss. bassin liouil. d. Valen., p. 502, pi lxxvii. fig. 1. 

Horizons and Localities. — 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Roof of New Mine Coal: Merry Hill Colliery, Brierley Hill. 
Roof of Bottom Coal: No. 120 Pit, Coneygre Colliery, Tipton. 

Sigillaria Brongniart. 
Sigillaria discophora Konig, sp. 

Lepidodendron discophorum, Konig, Icones foss. seciiles, pi. xvi. fig. 194. 

1885. Sigillaria discophora, Kidston, Ann. and Mag. Nat. Hist., 5th ser., vol. xvi. p. 251, pi. iv. fig. 5, 

pi. v. fig. 8, pi. vii. figs. 12, 13. 
1889. ,, ,, Kidston, ibid., 6th ser., vol. iv. p. 61, pi. vi. fig. 1. 

1831. Ulodendron minus, L, & H., Fossil Flora, vol. i. pi. vi. 

1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 483, pi. lxxiii. fig. 2, pi. lxxiv. 

%• 5. 
1831. Ulodendron majus, L. & H., Fossil Flora, vol. i. pi. v. 

Horizon and Locality. — Roof of Bottom Coal: No. 120 Coneygre Pit, Tipton. 
Sigillaria semipulvinata Kidston. 

1897. Sigillaria semipulvinata, Kidston, Trans. Roy. Soc. Edin., vol. xxxix. p. 57, pi. iii. figs. 1-5. 

Note. — The only specimen I have seen from the South Staffordshire Coal Field is 
the small example figured in the Trans. Roy. Soc. Edin., vol. xxxix. p. 58, pi. iii. fig. 4, 
which I received from the late Mr C. Bbale in 1887. 

Horizon and Locality. — Westphalian Series : Great Bridge, near Dudley. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 139 

Sigillaria elegans (Sternberg, sp. ?) Brongt. 

1826. (?) Favularia elegans, Sternb., Essai flore monde prim., vol. i. fasc. iv. pp. xiv and 48, pi. lii. fig. 4. 

1828. Sigillaria elegans, Brongt., Prodrome, p. 65. 

1836. ,, „ Brongt., Hist. d. veget.foss., p. 438, pi. cxlvi. fig. 1, pi. civ., pi. clviii. fig. 1. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 582, pi. lxxxvii. figs. 1-4. 
1911. ,, ,, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 185. 
1836. Sigillaria hexagona, Brongt., ibid., pi. civ., pi. clviii. fig. 1. 

1836. Sigillaria minima, Brongt., ibid., p. 43?, pi. lviii. fig. 2. 

1887. Sigillaria elegantula, Weiss, Sigillarien d. preuss. Steink., I. Gruppe Favularien, p. 44, pi. xiii. 

figs. 74-78. 

Horizons and Localities. — 

Roof of New Mine Coal : Doulton's Clay Pit, Netherton ; Mount Pleasant, 

Brierley Hill. 
Roof of Bottom Coal: No. 120 Pit, Coney gre Colliery, Tipton. 

Cf. Sigillaria nodosa Bowman, sp. 

PL XII. figs. 1, la. 

1836. Favularia nodosa, Bowman, in L. & H., Fossil Flora, vol. iii. p. 107, pi. cxcii. figs, a, b, c. 

Remarks. — It is probably impossible to ever ascertain the true nature of the plant 
which formed the type of this species. Of the three figures given by Lindley and 
Hutton on their plate, only that at a demands consideration, b and c being too 
imperfectly preserved, and even here one cannot help suspecting that their figure a is 
somewhat " restored." 

Among the specimens collected by Mr Hughes were some examples of a Sigillaria 
which from their state of preservation may throw some light on the nature of this 
interesting but not understood " species " — the Sigillaria nodosa Bowman. 

One of these specimens is shown natural size on PI. XII. fig. 1. The leaf scars are 
much flattened, and it requires careful examination to make out their form. This, 
however, can be done, and some of them are given enlarged at fig. la. 

The full description of this specimen is : — Stem ribbed, furrows slightly flexuous. 
Leaf scars approximate, divided by transverse furrows and occupying about § of the 
width of the rib, upper margin slightly emarginate, lower margin almost straight, 
lateral margins convex, with feebly developed lateral angles placed about the centre of 
the sides. Cicatricules three, situated about § above base of scar, the two lateral 
punctiform, the central punctate or slightly elongated transversely. 

The specimen has a great similarity to the Sigillaria dolearis Weiss, # but Koehne, 
when speaking of this type, says that from its state of preservation it cannot be 
determined whether it belongs to Sigillaria elegantula Weiss ( = Sigillaria elegans 
Brongt.) or to Sigillaria cumulata Weiss. Certainly our specimens are not the latter 
plant, but I am inclined to think they may be some form of Sigillaria elegans 
* Sigillarien d. preuss. Steink., I. Gruppe d. Favularien, p. 31, pi. iii. (ix.) fig. 37. 



140 DR ROBERT KIDSTON ON THE 

( = Sigillaria elegantula Weiss). But whatever species our fossils may belong to, 
their state of preservation has produced a fossil which might well be the Sigillaria 
nodosa Bowman. It is highly probable that Sigillaria nodosa has been founded on 
similar specimens to these ; but beyond a "belief" that this is the case, one cannot go 
further, for, unless the original specimens were examined, it is impossible to ascertain 
how much of the figure belongs to the specimen and how much to the artist. 

Taking, therefore, all the available data ink) consideration, it seems highly probable 
that Sigilla.ria nodosa Bowman, sp., has been founded on an imperfectly preserved 
specimen of one of the many forms of Sigillaria elegans Brongt. ( = Sigillaria 
elegantula. Weiss). 

I at one time thought that Sigillaria nodosa Bowman, sp., was a form of Sigillaria 
tessellata, Brongt., and a Sigillaria which is not very uncommon in the Eadstock Series 
I figured under the name of Sigillaria tessellata, var. nodosa ; # but this 1 now identify 
as the Sigillaria cumulata Weiss, and as far as my experience goes this species is 
restricted to the Eadstockian Series in Britain, whereas Sigillaria, nodosa Bowman 
originates from the Westphalian Series. 

Horizon and Locality. — Roof of New Mine Coal: Merryhill Colliery, Mount 
Pleasant, Brierley Hill. 

Sigillaria trigona Sternberg, sp. 

PI. XII. figs. 2, 2a, 3, and 4. 

1820. Lepidodendron irigonum, Sternb. (non Knorr), Essai flore rnonde prim., i., fasc. i. p. 25, pi. xi. 

fig. I- 
1826. Favularia trigona, Sternb., Essai flore monde prim., i., fasc, iv. p. xiii. 

1887. Sigillaria trigona, Weiss, Sigillarien d. preuss. Steink., part i. pp. 36 and 53, pi. v. fig. 54, pi. ix. 

(xv.) fig. 1 (Favularia trigona copied from Sternberg. Abhandl. d. geol. 
Landesanstalt, vol. vii. Heft iii.). 
1894. ,, ,, Kidston, Trans. M'ter. Geol. Soc, part xxi. vol. xxi. p. 642, text-fig. 

1893. Sigillaria subornata, "Weiss, Sigillarien d. preuss. Steink. u. Rothl. Gebiete: II. Subsigillarien, 

p. 209, pi. xxvii. fig. 106. 
1893. (?) Sigillaria decorala, Weiss, ibid., p. 207, pi. xxvii. fig. 105. 
1904. Sigillaria limbata, Zalessky, Mem. Comite geol., nouv. se>., livr. 13, pp. 74 and 122, pi. xiii. fig. 11. 

Description. — Stem ribbed, furrows strongly zigzag, leaf scars alternate, more or 
less close, pyriform, occupying almost the whole width of the rib and sitting on 
bracket-like elevations so that the basal margin holds a higher level than the apex and 
placed towards the upper end of subhexagonal areas of the rib, which are separated 
from each other by a straight or curved transverse furrow ; upper portion of leaf scar 
narrow, rounded, with concave sides and surmounted by a ligule pit, lower portion of 
scar wider, more obtusely rounded ; lateral angles prominent and situated about the 
centre of the scar. Cicatricules three, the central transversely semilunar and placed in 
the centre of the lateral cicatricules. A short ridge extends from the lateral angles to 
the furrows. Areas on which the leaf scars are placed are smooth, except for the 

* Proc. York. Geol. and Polytech. Soc, vol. xiv. pp. 353 and 385, pi. lviii. fig. 1, 1902. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 141 

presence of two short diverging ridges which descend from the base of the elevation 
of the leaf scar. 

The subepidermal surface of the ribs is longitudinally striated. 

Remarks. —Although the coaly matter has been removed from part of the specimen 
figured here, and partially fractured at other parts, still many of the leaf scars are 
beautifully preserved, and show very distinctly the details of their structure. 

Sigillaria trigona Sternb., with its prominent zigzag furrows, belongs to the Favu- 
laria section of the genus. The pyriform, or what Weiss has well described as " bell- 
shaped,"' leaf scars, contracted at top, occupy the narrower, and the expanded base the 
wider, portion of the rib. Bach leaf scar is separated from its neighbour above and 
below by a transverse furrow which passes across the whole width of the rib. These 
transverse furrows are sometimes straight as at fig. 3, or semicircular as at fig. 4, both 
forms occurring on the same specimen as seen on the enlarged portion given at fig. 2<x. 
The leaf scars are therefore placed in compartments which gradually slope upwards 
from the top to the base, and thus come to be situated on little sloping brackets. The 
line following the basal contour of the leaf scars seen on figs. 3 and 4 represents the 
elevation of the lower margin of the scar ; the cushion on which it sits is smooth, and 
free from all ornamentation except for the occurrence of two small ridges which are so 
slight that they can only be observed on well-preserved examples. 

Like many of the older species, owing to imperfect description some doubt has 
arisen as to the specific position of Sigillaria trigona Sternb. (non Knorr). The original 
figure shows the specimen to belong distinctly to the Favularia section of Sigillaria, and 
has " bell-shaped" leaf scars rounded below and somewhat truncate or rounded above. 

Though the upper margins of the leaf scars are mostly truncate on Sternberg's 
figure, a few are rounded at top like those shown on the figure given by Weiss # 
and that shown here. It appears to me therefore that these two specimens, and that 
figured some years ago under the same name by myself,f from the form of their 
leaf scars, the zigzag furrows, and the raised- up bracket-like shelf that supports the 
leaf scar, as well as their smooth ribs, refer them without doubt to the Sigillaria 
trigona Sternb. 

Sigillaria subornata Weiss should, I think, be united to Sigillaria trigona Sternb. 
The differences are so slight — the leaf scar a little broader in proportion to its length 
and the lunate part of the cushion below the scar not so wide ; but these differences can 
scarcely be considered as of specific value. It seems also highly probable that 
Sigillaria decorata Weiss belongs to the same plant. The occasional occurrence of a 
few transverse wrinkles on the cushion below the leaf scar, and the slightly greater 
distance of the leaf scars apart, are surely not sufficient characters for the creation of a 
"species," even when we add that the leaf scars are a little broader in proportion to 
their length. 

The specimens assigned to Sigillaria trigona by Feistmantel appear to have been 

* hoc. cit. t hoc. cit. 



142 DR ROBERT KIDSTON ON THE 

badly preserved, and possibly do not belong to this species. They are therefore 
excluded from the references.* From typical Sigillaria mamillaris Brongt., Sigillaria 
trigona is essentially distinct, the former belonging to the Rhytidol&pis section (those 
with straight or only slightly flexuous furrows), while the latter belongs to the 
Favularia section, but in Sigillaria Dournaisii Brongt., + which is now united to 
Sigillaria mamillaris as a variety of that species, when the scars are large and occupy 
almost the whole width of the ribs, which then become slightly enlarged and somewhat 
zigzag, some similarity between the two species occurs, but the " bell-shaped" scars of 
Siyillaria trigona and the pronounced zigzag furrows will at once distinguish them, 
and further, the rib below the leaf scar is always distinctly ornamented with transverse 
bars in Sigillaria mamillaris and its varieties. 

Still more closely related to Sigillaria trigona is the Sigillaria Hauchecornei 
Weiss.J Here, however, the leaf scars are slighly broader in proportion to their length 
and the two ridges below the leaf scars extend downward from the margin of the scar 
itself, while in Sigillaria trigona they are only on the cushion proper below the 
leaf scar.§ 

The Sigillaria trigona Sternb., sp., is very rare in Britain. 

Horizon and Locality. — Roof of Thick Coal : Tipton. 



Sigillaria tessellata Brongt. 

1818. Phgtolithus tessellatus, Steinhauer, Trans. Amer. Phil. Soc, vol. i. p. 295, pi. vii. fig. 2. 
1836. Sigillaria tessellata, Brongt., Hist. d. veget.foss., p. 436 ("? pi. clvi. fig. 1), pi. clxii. figs. 1-4. 
1886. „ „ Zeiller, Flore foss. bassin houil. d. Valen., p. 561, pi. Ixxxv. figs. 1-9, 

pl. lxxxvi. figs. 1-6. 
1911. ,, ,, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 188. 

1836. Sigillaria Knorrii, Brongt., ibid., p. 444, pl. clvi. figs. 2-3 (? pl. clxii. fig. 6). 
1836. Sigillaria alveolaris, Brongt. (non Sternb.), ibid., p. 443, pl. clxii. fig. 5. 
1848. Sigillaria contigua, Sanveur, Veget.foss. d. terr. houil. Belgique, pl. lii. fig. 1. 

Note. — Frequent. 
Horizons and Localities. — 

Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr. 
near Birmingham. 

Ten-foot Ironstone Measures : Cabbage Hall Pits, Netherton, near Dudley. 

Blue Measures : Doulton's Clay Pit, Netherton, near Dudley. 

Roof of New Mine Coal: Mount Pleasant, Brierley Hill. 

Roof of Fireclay Coal: Doulton's Clay Pit, Netherton. 

Roof of Bottom Coal: No. 120 Pit, Coneygre Colliery, Tipton. 

* Vers. d. bohm. Ahlager., Abth. iii. pl. ix. figs. 3-5, 1876. 

t Sigillaria Dournaisii Brongt., Hist. d. veget.foss., p. 441, pl. cliii. fig. 5. 

X Sigillaria d. preuss. Steink, No. I. p. 47, pl. xiii. figs. 81-82, 1887. 

§ Koehnk uniie.s Sigillaria Hauchecornei Weiss with Sigillaria mamillaris as a variety. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 143 

Sigillaria mamillaris Brongt. 

1824. Sigillaria mamillaris, Brongt., Ann. d. Scienc. nat., vol. iv. p. 33, pi. ii. fig. 5. 

1836. ,, ,, Brongt., Hist. d. veget.foss., p. 451, pi. cxlix. fig. 1 (1 pi. clxiii. fig. 1 var.) 

1886. ,, „ Zeiller, Flore foss. bassin houil. d. Valen., p. 577, pi. lxxxvii. figs. 5-10. 

1911. ., ,, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 190. 

1836. Sigillaria Dournaisii, Brongt., Hist. d. veget. foss., p. 441, pi. cliii. fig. 5. 

1876. Sigillaria conferta, Boulay, Terr, houil. du Nord de la France, p. 44, pi. iii. fig. 3. 

Horizons and Localities. — 

Blue Measures, six feet above BroocJi Coal : Hamstead Colliery, Great Barr, 

near Birmingham. 
Roof of Brooch Coal : Shut End. 
Roof of Thick Coal: Bradley Colliery, Bilston. 

Roof of Fireclay Coal: Doulton's Clay Pit, Netherton, near Dudley. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 



forma abbreviata Weiss. 

1871. Sigillaria mamillaris /3 abbreviata, Weiss, Fossil Flora d. jiingst. Stk. u. Roihl., p. 165, pi. xv. 
figs. 1, 2. 

Horizon and Locality. — Whitestone Measures: Moxley, near Bilston. Collected 
by the late Mr C. Beale. (K. No. 838.) 



forma Dournaisi Brongt., pro. sp. 

Horizon and Locality. — Roof of Fireclay Coal: Doulton's Clay Pit, Netherton, 
near Dudley. 

Sigillaria scutellata Brongt. 

1822. Sigillaria scutellata, Brongt., Class, d. veget. foss., pp. 22 and 89, pi. i. fig. 4. 

1836. „ ,, Brongt., Hist. d. veget.foss., p. 455, pi. cl. figs. 2, 3, pi. clxiii. fig. 3. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 533, pi. lxxxii. figs. 1-6, 9. 

1911. „ ,, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, p. 192. 

1836. Sigillaria notata, Brongt., ibid., p. 449, pi. cliii. fig. 1. 

1836. Sigillaria elliptica, var., Brongt., ibid., p. 447, pi. clxiii. fig. 4. 

Horizon and Locality. — Roof of New Mine Coal : Merryhill Colliery, Brierley Hill. 

Sigillaria vulgaris Artis, sp. 

1825. Euphorbites vulgaris, Artis, Antedil. Phyt., pi. xv. 

Remarks. — Formerly I treated this species as a variety of Sigillaria scutellata 
Brongt.,* but in more recent years I have seen a number of specimens from the 

* Catal. Palmoz. Plants, p. 186, 1886. 



144 DR ROBERT KIDSTON ON THE 

Westphalian Series which are characterised by the " bell-shaped " leaf scars, as figured 
by Artis. It is therefore here provisionally treated as a distinct species. 

The specimen recorded from the South Staffordshire Coal Field is in the collection 
of Mr H. W. Hughes, and very typical of the plant described by Artis as Euphorbites 
vulgaris. I find the distance apart of the leaf scars varies considerably on different 
specimens, on some being approximated and having much the appearance of Sigillaria 
polita Lesqx.* Lesquereux describes his specimen as " ribs flat, very smooth," but 
on the better-preserved specimens of Sigillaria vulgaris in my collection the lower 
margin of the leaf scar is elevated, and the descending lines from the low-placed lateral 
angles have transverse markings as in Sigillaria mamillaris, to which perhaps it has 
a closer affinity than to Sigillaria scutellata. 

Koehne unites the Euphorbites vulgaris provisionally with Sigillaria Walchi.f 
Horizon and Locality. — Old Mine Clay = Stinking Coal : Docking-iron Pit, Delph, 
Brierley Hill. 

Sigillaria cordigera Zeiller. 

PI. XII. figs. 5, 5a. 

1886. Sigillaria cordigera, Zeiller, Flore foss. basein liouil. d. Valen., p. 526, pi. Ixxviii. fig. 5. 

Description. — Stem ribbed, ribs slightly arched, separated by well-marked straight 
furrows. Surface of ribs ornamented with a fine shagreen, especially prominent between 
the leaf scars, where fine short transverse line-like little furrows break up the surface 
of the cortex. Leaf scars occupy about half width of rib, cordate, rounded, and 
slightly contracted at base, with a large sinus on the upper margin, distant from 
each other about the length of a leaf scar. Lateral angles absent. Cicatricules placed 
about the distance of ^ the length of the leaf scar from the top, central (vascular) 
cicatricule punctiform or slightly transversely elongated, the two lateral (parichnos) 
oval, vertical or slightly diverging and on the same level as the vascular cicatrice 
Ligule pit situated immediately above the leaf scar. 

Remarks. — This species, which appears to be very rare on the Continent, has not 
been previously recorded for Britain, and is only known from a single locality. 

Sigillaria cordigera is at once distinguished from all the other species of the 
genus by its cordate leaf scars, which frequently show a slight irregularity in form. 

At fig. ba a small part of the specimen is enlarged 2 times to show the 
ornamentation of the cortex, which has much the appearance of the shagreen of 
leather. 

I am indebted to Mr H. W. Hughes for a specimen of this species which was 
collected by himself. 

Horizon and Locality. — Roof of New Mine Coal: Mount Pleasant, Brierley Hill. 

* In Rogers, Geol. of Pennsyl, p. 872, pi. xiv. fig. 3, 1858. Goal Flora, p. 490, pi. lxxiii. fig. 1. 

t <: Sigillariastamme," Abhandl. d. konigl. preuss. geol. Landesanstalt, neue Folge, Heft xliii. p. 54, 1904. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 145 

Sigillaria reniformis Brongt. 

1824. Sigillaria reniformis, Brongt., Ann. d. science nat., vol. iv. p. 32, pi. ii. fig. 2. 

1836. ,, ,, Brongt., Hist. d. veget. foss., p. 470, pi. cxlii. 

1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 556, pi. lxxxiv. figs. 4-6. 

1888. ,, ,, Kidston, Trans. Roy. Soc. Edin., vol. xxxv. p. 327, plate, fig. 2. 

1911. ,, ,, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 194. 

1848. Sigillaria grandis, Sauveur, Veget. foss. d. terr. houil. de Belgique, pi. lvii. fig. 1. 

1876. Sigillaria laticostata, Boulay, Terr, houil. du Nord. de la France, p. 46, pi. iii. fig. 2. 

Horizon and Locality. — Blue Measures, above Brooch Coal : Hamstead Colliery, 
Great Barr, near Birmingham. 

Sigillaria elongata Brongt. 

1824. Sigillaria elongata, Brongt., Ann. d. science nat., vol. iv. p. 33, pi. ii. figs. 3, 4. 
1836. ,, ,. Brongt., Hist. d. veget. foss., p. 473, pi. cxlv., pi. cxlvi. fig. 2. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 545, pi. lxxxi. figs. 1-9. 

1911. ,, ,, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 202. 

1836. Sigillaria Cortei, Brongt., ibid., p. 467, pi. cxlvii. figs. 3, 4. 
1836. Sigillaria intermedia, Brongt., ibid., p. 474, pi. clxv. fig. 1. 
1836. Sigillaria grseseri, Brongt., ibid., p. 454, pi. clxiv. fig. 1. 
1836. Sigillaria gracilis, Brongt., ibid., p. 462, pi. clxiv. fig. 2. 

1888. Sigillaria mamillaris Kidston (wore Brongt.), Trans. Roy. Soc. Edin., vol. xxxv. p. 328, plate, 
fig. 10. 

Remarks. — The specimen from Hamstead, which I identified as Sigillaj'ia mamillaris, 
and is figured in the Trans. Roy. Soc. Edin., vol. xxxv., plate, fig. 10, I now believe to 
be a form of Sigillaria elongata Brongt., with unusually close leaf scars. 

Horizon and Locality. — Blue Measures, above Brooch Coal: Hamstead Colliery, 
Great Barr, near Birmingham. 

Sigillaria rugosa Brongt. 

1836. Sigillaria rugosa, Brongt., Hist. d. veget. foss., p. 476, pi. cxliv. fig. 2. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 551, pi. lxxx. figs. 1-5. 

1911. ,, ,, Kidston, Mem. Musee roy. d'hist. nat. de Belgique, vol. iv. p. 203. 

Horizon and Locality. — Roof of Bottom Coal: No. 120 Pit, Coneygre Colliery, 
Tipton. 

Collected by Mr H. W. Hughes, F.G.S. 

Sigillaria punctirugosa Kidston, n. sp. 
PL XII. figs. 6-6a. 

Description. — Stem ribbed, furrows straight, leaf scars distant, occupying the 
whole width of rib, rounded-pentagonal, about as broad as high, slightly narrowing 
above the low-placed blunt lateral angles, straight or very feebly emarginate on upper 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 19 



146 DR ROBERT KIDSTON ON THE 

margin, lower margin rounded. Cicatricules three, placed about a third of length of leaf 
scar from top — the two lateral elongate, diverging ; the central punctiform, and on level 
with centre of lateral cicatricules. Ligule pit immediately above leaf scar, a short 
distance above which is a straight transverse furrow. Surface of rib ornamented with 
a leather-like shagreen and two slightly diverging rows of prominent transversely 
elongated little elevations which descend from the base of the scars and terminate at 
the transverse furrow. 

Remarks. — The position of the blunt lateral angles is below the centre of the leaf 
scar ; and these being very obscure, give it a rounded appearance. 

The shagreen ornamentation of the cortex, of which the reticulations are relatively 
large, and the two descending rows of elongated elevations, give a distinctive character 
to this species. 

Sigillavia punctirugosa is very rare, and has only been met with at one locality by 
Mr H. W. Hughes. 

Horizon and Locality. — Roof of Neiv Mine Coal: Mount Pleasant, Brierley Hill. 

Sigillariostrobus Schimper. 
Sigillariostrobus ciliatus Kidston. 

1897. Sigillariostrobus ciliatus, Kidston, Trans. Roy. Soc. Edin., vol. xxxix. p. 53, pi. ii. figs. 2-9. 

Remarks. — A small slab received from Mr Henry Insley contains some isolated 
but characteristic bracts of this species. 

Horizon and Locality. — Blue Measures, six feet above Brooch Coal : Hamstead 
Colliery, Great Barr, near Birmingham. 

Sigillariostrobus sp. 

Remarks. — Included here is a part of a cone which is closely allied to Sigillario- 
strobus Goldenbergi Feistmantel,* but it is slightly smaller, and the bracts are ciliate. 

Horizon -and Locality. — Ten-foot Lronstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

Collected by Mr H. W. Hughes. 

Cyperites L. & H. 
Cyperites bicarinata L. & H. 

1832. Cyperites bicarinata, L. & H., Fossil Flora, vol. i. pi. xliii. figs. 1, 2. 

Note. — This species is probably founded on fragments of Sigillarian leaves. 
Horizon and Locality. — Blue Measures, above Brooch Coal: Hamstead Colliery, 
Great Barr, near Birmingham. 

* See Zeiller, Flore foss. bassin houil. d. Valen., p. 600, pi. lxxxix. figs. 1 and 4. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 147 

Stigmaria Brongniart. 
Stigmaria ficoides Sternberg, sp. 

1820. Variolaria ficoides, Sternb., Essai flore monde prim., vol. i. fasc. i. pp. 23 and 26, pi. xii. figs. 1-3. 
1822. Stigmaria ficoides, Brongt., Class d. veget. foss., pp. 28 and 89, pi. i. fig. 7. 
1911. „ „ Kidston, Mem. Musee roy. d'hist. nat. de Belgique, p. 212. 

Note. — Generally distributed throughout the whole series. 

Stigmaria reticulata Gopp. 

1841. Stigmaria ficoides, var. reticulata, Gopp., Gatt. d. foss. Pflanzen, Lief. 1, 2, p. 30, pi. ix. fig. 11. 
1886. ,, ,, ,, Zeiller, Flore foss. oassin houil. d. Valen., p. 613, pi. xci. fig. 5. 

1862. Stigmaria anabathra, var. reticulata, Goldenberg, Flora Sareepont. foss., Heft iii. p. 19, pi. xiii. 

fig. 15. 
1836. Stigmaria ficoides, Gopp., Syst. fil. foss., p. 92, pi. xxxiii. fig. 7. 

Note. — This fossil has usually been treated as a variety of Stigmaria ficoides, and 
as such it was described by Goppert, but its distinctive characters fully entitle it to 
specific rank. 

Horizons and Localities. — Roof of Fireclay Coal: Russell's Hall, Dudley; Doulton's 
Clay Pit, Netherton. 

Lepidocarpon Scott. 

1900. Lepidocarpon, Scott, Proc. Roy. Soc. London, vol. lxvii. p. 306. 

1901. „ Scott, Phil. Trans., B, vol. cxciv. p. 325. 

Before giving a description of the new Lepidocarpon from Coseley, it is necessary 
to keep clearly before us the characters of this genus, and this can be best accomplished 
by transcribing here the diagnosis given by Dr Scott in his paper on " The Seedlike 
Fructification of Lepidocarpon, a genus of Lycopodiaceous Cones from the Carboniferous 
Formation " : — # 

" Strobili, with the characters of Lepidostrobus, but each megasporangium 
enclosed, when mature, in an integument growing up from the superior face of the 
sporophyll-pedicel. 

" Integument, together with the lamina of the sporophyll, completely enveloping 
the megasporangium, or nucellus, leaving only an elongated, slit-like micropyle above. 
A single functional megaspore or embryo-sac developed in each megasporangium and 
occupying almost the whole of its cavity. 

" Megaspore ultimately filled by the prothallus or endosperm. Sporophyll, together 
with the integumented megasporangium and its contents, detached entire from the axis 
of the strobilus, the whole forming a closed, seed-like, reproductive body. 

" Seed-like organ horizontally elongated, in the direction of the sporophyll-pedicel, 
to which the micropylar crevice is parallel." 

* Phil. Trans., loc. cit., p.^325. 



148 DR ROBERT KIDSTON ON THE 

To illustrate the character of Lepidocarpon more clearly, a diagrammatic repre- 
sentation of a sporophyll as seen in transverse section — that is, tangential to the cone 
axis — is given at text-fig. 7. Here the only one of the four megaspores in each 
sporangium which becomes developed is seen at mg, enclosed within the sporangium 
sp, which is attached to the sporophyll at b and enclosed by the integument i. This 
envelops the sporangium on both sides, only leaving at the summit a narrow slit m, 
which passes radially along the whole length of the elongated sporangium. 

m. 




m 



5 



Text-fig. 7. — Lepidocarpon Lomaxi Scott. Diagrammatic section tangential to axis. Description in text. 

In the fossil about to be described, as only the remains of some of these structures 
are preserved, their interpretation- is made much clearer by reference to this text-fig. 
The same distinguishing letters are used on the plate for the corresponding structures 
shown in the fossil. 

Lepidocarpon Westphalicum Kidston, n. sp. 
Plate XIII. figs. 1-6. 

Description. — Cone of unknown length, l - 40 cm. in diameter; sporophylls forming 
steep spirals and placed at right angles to axis ; pedicel on which sporangium sits 
5 "5 mm. long; sporangium oblong, compressed, with a basal keel, truncate at distal 
extremity, slightly contracted and rounded at proximal end, 5*5 mm. long, about 
2*5 mm. high, and slightly over 1 mm. broad ; integumented. On removal from cone 
the sporangiophores leave a narrow rhomboidal scar with central point on the axis. 

Remarks. — This remarkable fossil occurs in an ironstone nodule about 3 "50 cm. long 
and 2 "80 cm. broad. Its mode of preservation is most peculiar. The greater part of 
the cone is absent, but a small portion towards the centre is preserved, and this stands 
up free from the surface of the stone entirely uncompressed. The whole of the bracts 
have disappeared, except perhaps their bases between the sporangia, and the only parts 
exhibited are the axis, the sporangia, and the remains of the integuments. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 149 

The cone is seen natural size at figs. 1 and 2. Fig. 1 shows a surface view of the 
fossil, and fig. 2 an end view to show the elevation of the cone above the surface of 
the stone. 

Several enlargements of the specimen are given, under different directions of 
lighting, to exhibit the structure as fully as possible. 

On Plate XIII. fig. 3 the specimen is enlarged 2 times to give a general idea of the 
fossil, for owing to its small size this is not well exhibited at fig. 1. At fig. 4 it is 
enlarged 4 times to bring out more clearly the details of structure. 

The sporangiophores have disappeared, unless some indistinct remains below the 
sporangia may represent them ; but the sporangia are very clearly seen, arranged round 
the axis in steep spirals. They are compressed laterally, but this is not the result of 
external pressure, as the cone is uncompressed. Between the sporangia are seen the 
remains of the integuments, whose ruptured margins are seen between the sporangia, 
fig. 6, i. The scars left on the axis by the removal of the bracts are fusiform, with 
a central point. They are well seen on the axis, immediately above and below the 
sporangia, on fig. 4, x. They have very much the character of an imperfectly preserved 
Lepidodendroid leaf scar. They are also seen on fig. 6. The sporangia are slightly 
narrowed towards the top, fig. 4, 5, 6, sp. A good side view of the sporangia is seen at 
fig. 5, sp. The light strikes the specimen here in a different direction from that 
employed in photographing fig. 4. The base of the sporangium is seen to have a slight 
keel, probably indicating the line along which it was attached to the sporophyll. It 
will be noticed that there is a slight crack at the distal end of this sporangium, 
fig. 5, sp, but this is evidently an accidental occurrence. Another of the sporangia has 
had a piece broken off, and one can see into the now empty cavity, and the fracture 
shows that the sporangium wall must have been of considerable thickness. Fig. 6, sp'. 

The central portion of the cone has been enlarged about 8 times at fig. 6, with 
the object of showing more clearly the remains of the integuments. The sporangia are 
seen to be arranged in steep spirals, in which only four sporangia in each spiral now 
remain. The cone had probably reached the condition of maturity, as the integuments 
of the sporangia seem to have been fully developed, and their remains are seen at the 
places lettered, as well as at other parts of the fossil. A fragment of an integument 
stands out at i, and the other part is seen on the left, % . A small piece of the integu- 
ment of the neighbouring sporangium is also preserved, and at several places the remains 
of the integuments of contiguous sporangia are seen as two delicate structures, more or 
less following the contour of the sporangia. This can be observed at several places, 
especially at fig. 6, i' and i". The presence of these structures shows that the cone is 
not an ordinary Lepidostrobus, because in that genus the sporangia are naked. On the 
other hand, the structures just mentioned are identical in position and structure, as far 
as they show, with the integument in Lepidocarpon . The thick wall of the sporangium, 
which is seen on fig. 6, sp, though much importance cannot be placed on this, is also a 
character of Lepidocarpon. There is still the further circumstance that the missing 



150 



DR ROBERT KIDSTON ON THE 



sporangia and sporophylls have been entirely removed. There are no fragments of bases 
attached to the axis, which probably would have been the case if they had been forcibly 
broken off. As the sporangia are perfect and are not mixed with broken individuals, 
one has some ground for believing that the missing sporophylls and attached sporangia 
have been shed naturally at maturity. 

Notwithstanding, then, the fragmentary nature of our fossil, the characters it shows, 
point, I think, beyond all doubt to its being a typical Lepidocarpon. 

The only species with which it is necessary to compare it is Lepidocarpon Lomaxi 
Scott.* From this the Staffordshire specimen is at once distinguished by the size and 
form of the sporangia, and a comparison of the characters of this single organ will at 
once show their specific individuality. 



Lepidocarpon Lomaxi Scott. 
Sporangium broad at the base and contracting 
upwards into a point : 

Maximum vertical height 4 - 5 mm. 
,, tangential width 2 '5 mm. 
,, radial length 6'7 mm. 
(Measurements from immature cone. Mature 
cones are larger.) 

Cone after development of integuments : 
3 cm. diameter at base. 
2 cm. diameter at top. 



Lepidocarpon Westphalicum Kidston. 
Sporangium narrow, of about equal width, and 
only very slightly narrower at its upper margin : 
Maximum vertical height 2 "5 mm. 
tangential width 10 mm. 
,, radial length 5 - 5. mm. 
(Measurements from cone apparently mature.) 

Cone after develojmient of integuments : 

1"40 or 1"50 cm. probably about centre of 
cone. 



The distinction between the two species is therefore very marked. Of the two 
previously described species, one, Lepidocarpon Wildianum Scott, t is derived from the 
Calciferous Sandstone Series (Culm), of Petty cur, Fife, and the other, Lepidocarpon 
Lomaxi, from the Lanarkian Series (Lower Coal Measures) of Lancashire and 
Yorkshire. 

Lepidocarpon Westphalicum, as indicated by the name I propose for the new 
species, comes from the Westphalian Series, and so increases our knowledge of the 
vertical distribution of the genus. ' 

The specimen was collected by Mr H. W. Hughes, F.G.S., by whom it was placed 
in my hands for description. 

Horizon and Locality. — Ten-foot Lronstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Cordaitese. 

Cordaites Unger. 

Cordaites borassifolius Sternb., sp. 

1832. Flabellaria borassifolia, Sternb., Kssai flore monde prim,, vol. i. fasc. ii. pp. 31 and 36, pi. xviii. ; 

fasc. iv. p. xxxiv. 
1845. ,, ., Conla (para), Beitr. z. Flora d. Voncelt, p. 44, pi. xxiv. figs. 1-3 and 8. 



* Phil. Trans., ser. B, vol. cxcix. p. 294, plates, 1901. 
t Ibid., p. 314, 1901. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 151 

1849. Pychnophyllumborassifolium, Brongt., Tableau d. genres d. viget. foss., p. 65. 

1850. Cordaites borassifolius, Unger, Genera et Species, p. 277. 

1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 625, pi. xcii. figs. 1-6. 

1910. ,, ,, Renier, Paleontologie, pi. cxii. 

Horizons and Localities. — 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Blue Measures, six feet above Fireclay Coal : Doulton's Clay Pit, Netherton, 

near Dudley. 
Between Fireclay Coal and Bottom Coal : Doulton's Clay Pit, Netherton. 



Cordaites principalis Germar, sp. 

1848. Flabellaria principalis, Germar, Vers. v. Wettin u. Lobejun, p. 55, pi. xxiii. 

1855. Cordaites principalis, Geinitz (pars), Vers. d. Steinkf. in Sachsen, p. 41, pi. xxi. figs. 1-2, 

2a, 2b. 
1886. ,, ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 629, pi. xciii. fig. 3, pi. xciv. 

fig- I- 
1911. ,, ,, Kidston, Mem. Musee roy. d'hist. nat. d. Belgique, p. 232. 

1833. Knorria taxina, L. & H., Fossil Flora, vol. ii. pi. xcv. a (stem). 

Horizons and Localities. — 

Blue Measures, above Brooch Coal: Jubilee Pit, Sandwell Park, West 

Bromwich ; Hamstead Colliery, Great Barr, near Birmingham. 
Roof of Brooch Coal: Holly Hall, Dudley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 
Roof of New Mine Coal : Clattershall Colliery, Brettell Lane. 
Blue Measures, six feet above Fireclay Coal : Doulton's Clay Pit, Netherton, 

near Dudley. 



Cordaites (Dory cordaites) palmseformis Gopp., sp. 

1852. Nmggeratliia palmxformis, Gopp., Foss. Flora d. Uebergangs, p. 216, pi. xv., pi. xvi. figs. 1-3. 

1855. ,, ,, Geinitz, Vers. d. Steinkf. in Sachsen, p. 42, pi. xxii. fig. 7. 

1864. ,, ,, Gopp., Foss. Flora d. perm. Form., p. 157, pi. xxi. fig. 26, pi. xxii. 

figs. 1-2. 
1871. Cordaites palmxformis, Weiss, Foss. Flora d. jiingst. Steink. u. Rothl., p. 199, pi. xviii. fig. 39. 

1876. „ „ Heer, Foss. Flora Helv., p. 56, pi. i. fig. 18. 

1877. Cordaites (Dorycordaites) palmxformis, Grand'Eury, Flore carbon, du depart, de la Loire, 

p. 214, pi. xviii. figs. 4, 5. 
1886. Dorycordaites palmxformis, Zeiller, Flore foss. bassin houil. d. Valen., p. 632, pi. xciii. 

figs. 1,2. " 
1890. Cf. Dorycordaites palmseformis, Renault, Flore foss. terr. houil. de Commentry, part ii. p. 585, 

pi. lxvi. figs. 1-7. 
1874. Pycnophyllum palmxforme, Schimper, Traite d. paleont. veget., vol. iii. p. 563. 

Horizon and Locality. — Blue Binds, above Brooch Coal : Jubilee Pit, Sandwell 
Park Colliery, West Bromwich. Collected by Mr H. W. Hughes. 



152 DR ROBERT KIDSTON ON THE 

Cordaites Brandlingi Witham, sp. 

1831. Witham, Observations on Fossil Vegetables, p. 31, pi. iv. figs. 1-6. 

1831. Pinites Brandlingi, L. k H., Fossil Flora, vol. i. pi. i. 

1833. „ ,, Witham, Internal Structure of Fossil Vegetables, pp. 43 and 73, pi. ix. figs. 

1-6, pi. x. figs. 1-6, pi. xvi. fig. 3. 

1S46. Araucarites Brandlingi, Gbpp., in Tchihatcheff, Voyage scient. dans V Altai Oriental, p. 389. 
1850. ,, ,, Grbpp., Foss. Coniferen, p. 232, pis. xxxix., xl., xli. figs. 1-7. 

1S48. ,, „ Germar, Vers. d. Stein/c. v. Wettin u. Lobejun, p. 49, pis. xxi.-xxii. 

1847. Dadoxylon Brandlingi, Endlicher, Syn. coniferarum fossilium, p. 35. 

1870. Araucarioxylon Brandlingi, Kraus, in Schimper, Traite d. paleont. veget., vol. ii. p. 382. 

1883. Araucaroxylon Brandlingi, K. Feistmantel, " Ueber Araucaroxylon in der Steink. von Mittel- 

Bbhmen," Abhandl. d. kbnigl. bbhm. Gesell. d. Wissen., iv. Folge, xii. Band, p. 17, pi. ii. 

figs, la, lb, lc, and 1a. 
1877. Dadoxylon (Cordaixylon) Brandlingi, Grand'Eury, Flore carb. du depart, de la Loire, p. 264. 
1890. Cordaioxylon Brandlingi, Schenk, in Zittel, Handbuch d. Palaeont., ii. Abth. Palaeophyt.,]). 853, 

fig. 408 (not fig. 173, p. 243). 
1888. Cordaites Brandlingi, Gbpp. and Sterzel, " Nachtrach. z. Kennt. d. Coniferenhblzer d. Palaeoz. 

Form.," Kbnigl. preuss. Akad. d. Wissensch., p. 12, pi. i. figs. 1-4. 
1900. „ ,, Penhallow, Trans. Roy. Soc. Canada, 2nd ser., vol. vi. p. 62, figs. 1, 9, 

10, 11. 
1900. Cordaites (Araucarioxylon) Brandlingi, Scott, Studies in Fossil Botany, p. 418, fig. 137 ; 2nd ed., 

p. 528, fig. 190, 1909. 

Note. — The only specimens of this species which 1 have seen from the South 
Staffordshire Coal Field were a few fragments collected by the late Mr C. Beale in 
1887. One of these is exceptionally well preserved, and is that from which the figure 
of Cordaites [Araucarioxylon) Brandlingi given by Scott (loc. cit.) was prepared. 

Horizon and Locality. — Westphalian Series : Rowley Regis, near Dudley. 

Artisia Sternberg. 
Artisia approximata Brongt. 

181*8. Phytolithus transversus, Steinhauer, Trans. Amer. Phil. Soc, p. 293, pi. v. fig. 3. 
1828. Sternbergia approximata Brongt., Prodrome, p. 137, 

1837. ,, ,, L. & H., Fossil Flora, vol. iii., pis. ccxxiv., ccxxv. 

1838. Artisia approximata, Corda, in Sternberg, Essaiflore monde prim., vol. ii. fasc. vii.-viii., p. xxii., 

pi. liii. figs. 1-6. 
1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 634, pi. xciv. figs. 2, 3. 

1890. ,, ,, Renault, Flore foss. terr. houil. d. Gommentry, part ii. p. 581, pi. Ixv. fig. 4. 

1893. „ „ Sterzel, "Flora d. Rothl. im Plauen. Grunde," p. 109, pi. x. fig. 4 (Abhandl. 

d. kbnigl. sdchsischen Gesell. d. Wissensch., vol. xxxii., 1893). 
1910. „ ,, Renier, Paleontologie, pi. cxvi. 

1848. Sternbergia transversa, Sauveur (non Artis), Veget. foss. d. terr. houil. Belgique, pi. lxix. fig. 1. 
1848. Sternbergia minor, Sauveur, ibid., pi. lxix. fig. 2. 

1869. Artisia transversa, Roehl (non Artis), Foss. Flora d. Steink.-Form. Westph., p. 148, pi. iv. fig. 8. 

1876. ,, ,, Roemer (non Artis), Letha>.a geog., vol. i. p. 242, pi. Iv. fig. 3. 

1886. Artisia sp., Sterzel, Flora d. Rothl. im Nordio. Sachsen. (Dames u. Kayser, Palaeont. Abliand., 

Band iii. Heft. 4), p. 32, and explan. to pi. iv. figs. 5, 6. 
1899. Artisia Steinkern., Potonid, Lehrb. d. Pflanzenpal., p. 267, fig. 253. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 153 

1901. Artisia Cor daites principalis, Stefani, Flore carbon, e perm, della Toscana, p. 103, pi. xiv. fig. 3. 
1860. Lomatophloios crassicaulis, Eichwald (pars), Lethxa Rossica, p. 156, pi. ix. fig. 3. 
1869. Lomaiofloios crassicaule, Roehl (pars), Foss. Flora d. Steink.-Form. Westph., p. 146, pi. xx. fig. 3, 
pi. xxiv. fig. 3. 

Horizons and Localities. — 

Roof of Brooch Coal : Himley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 

Oordaianthus Grand'Eury. 
Oordaianthus Volkmanni Ett., sp. 

1852. Catamites Volkmanni, Ett. (pars), SteinJcf. v. Stradonitz, p. 5, pi. v. figs. 1-3 (non fig. 4, non 

pi. vi. figs. 1, 2). 
1886. Cordaianthus Volkmanni, Zeiller, Flore foss. bassin houil. d. Valen., p. 637, pi. xciv. fig. 6. 
1874. Antholithus parviflorus, Schimper, Traite d. paleont. veget., vol. iii. p. 567, pi. ex. figs. 1-3. 

Note. — Rare. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. Collected by Mr H. W. Hughes. 

Oordaianthus Pitcairnise L. & H., sp. 

1833. Antholithus Pitcairnise, L. & H., Fossil Flora, vol. ii. pi. lxxxii. 

1881. Cordaianthus Pitcairnise,, Renault, Cours. d. bot.foss., vol. i. p. 94, pi. xiii. fig. 7. 

1886. „ ,, Zeiller, Flore foss. bassin houil. d. Valen., p. 639, pi. xciv. figs. 4, 5. 

1911. ., ,, Kidston, Mem. Musee roy. d'hist. not. de Belgique, vol. iv. p. 235. 

1910. ,, „ Renier, Paleontologie, pi. cxiii. 

1872. Cardiocarpon Lindleyi, Carr, Geol. Mag., vol. ix. p. 55, figs. 1, 2. 

Horizons and Localities. — 

Blue Measures, six feet above Brooch Coed : Jubilee Pit, Sandwell Park 

Colliery, West Bromwich. 
Roof of Brooch Coal : Shut End. 

Cordaianthus sp. 

Remarks. — A form allied to Cordaianthus Pitcairnise, but apparently specifically 
distinct from that species. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley 

Cordaianthus sp. 

Several specimens of a form of Cordaianthus allied to Cordaianthus major 
Renault * have been collected at Sandwell by Mr H. W. Hughes. 

The specimens bear oval seeds, distichously placed, from 1 '30 cm. to 1*90 cm. long, 

* Flore foss. terr. houil. d. Gommentry, part ii. p. 593, pi. lxxii. figs. 33, 34, 1890. 
TRANS. ROY. SOC. EDIN, VOL. L. PART I. (NO. 5). 20 



154 DR ROBERT KIDSTON ON THE 

according to their position on the axis. Occasionally the seeds appear as if enclosed in 
bracts, and at some places a large subtending bract is present. 

These are the first British specimens of the kind which I have met with. 

Horizon and Locality. — Blue Measures, above Brooch Coal: Jubilee Pit, Sand- 
well Park Colliery, West Bromwich. 



SEEDS. 

Samaropsis Goppert. 

Samaropsis Gutbieri Geinitz, sp. 
PI. XVI. fig. 2. 

1855. Cardiocarpon Gutbieri, Geinitz, Vers. d. Stemkf. in Sachsen, p. 39, pi. xxi. figs. 23-25. 
1888. Cardiocarpus Gutbieri, Kidston, Trans. Roy. Soc. Edin., vol. xxxiii. p. 403, pi. xxiii. fig. 5. 
1893. ,, ,, Potonie^ Flora d. Rothl. v. Thiiringen, p. 254, pi. xxxi. figs. 15-19. 

1881. Cardiocarpus sclerolesta, Brongt., and Cardiocarpus sclerotesta minor, Brongt., Reciter, graines foss. 
silic, p. 37, pi. A, figs. 5, 6. 

Description. — Winged cordate seeds longer than broad or broader than long, with 
a sharp point and a rounded or notched base. Wing rather narrow, wider at base, and 
gradually becoming narrower as it reaches apex. Nut lenticular in section. 

Remarks. — These seeds vary much in size and in the relative proportions of length 
to width. According to Geinitz they attain a width of 2 cm., the same size as that 
figured here ; but Potonie, who unites Cardiocarpon reniforme Geinitz # with Samar- 
opsis Gutbieri, ascribes to them a width of 3-5 cm. 

Potonie has suggested that the specimens of Cardiocarpus major figured by 
Grand'Eury t and Renault J may be isolated nucules of Samaropsis Gutbieri. This 
is most probably the case, for in the small specimen I figured from the Radstock 
Coal Field § the nucule lifts out of the matrix, and in form is identical with the 
Cardiocarpus major, as figured by these authors, only slightly smaller. In my 
specimen, however, the nucule when replaced in the little hollow in the matrix is seen 
to have been surrounded by a wing which occurs as a stain on the stone. 

Horizons and Localities. — 

Shale over Brooch Coal : Holly Hall, near Dudley. 

Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 

Roof of Thick Coal : Tipton. 

Samaropsis Meachemi Kidston, sp. 

1888. Cardiocarpus Meachemi, Kidston, Trans. Roy. Soc. Edin., vol. xxxv. p. 330, plate, figs. 5-7. 

Horizon and Locality. — Blue Measures, above Brooch Coal : Hamstead Colliery, 
Great Barr, near Birmingham. 

* Dyas, p. 145, pi. xxxi. fig. 16, 1861-62. t Flore carbon, du depart, de la Loire, p. 235, pi. xxvi. fig. 16. 

X Flore foss. terr. houil. d. Commentry, part ii. p. 600, pi. lxxii. figs. 10, 11, 1890. § hoc. cit. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 155 

« 

Samaropsis quadriovata Kidston, n. sp. 
PI. XVI. figs. 1, la. 

Description.— Seed winged, quadrate-oval, 9 mm. long and 7 mm. wide. Nucule 
cordate-acute, with a slight basal notch 6 mm. long and 5 '50 mm. wide at base ; wing 
very narrow at base, but gradually widening as it advances upwards towards the apex, 
where it is slightly emarginate and attains a width of fully 2 mm., for about which 
distance it extends beyond the apex of the nucleus. 

Remarks. — The seed is shown natural size on PI. XVI. fig. 1 and enlarged 2 
times at fig. la. It is somewhat of the type of Samaropsis fiuitans Dawson,* but is 
larger, more quadrate, and the wing is not so much developed at the upper end of 
the seed. 

There are several other described species of Samaropsis to which Samaropsis 
quadriovata is more or less closely related, but with the figures and descriptions of 
these there is none with which it accurately agrees. Among these allies may be 
mentioned Cardiocarpus simplex Lesqx.,t but this has a wing of about equal width all 
round the nucule. To Cardiocarpus (Samaropsis) latealatus Lesqx. \ it has also some 
similarity, but this also has a wider wing at the base and is more contracted into a 
point at the apex. 

With one of the figures of Cardiocarpus crassus Lesqx., that given in the Coal 
Flora, vol. iii. p. 812, pi. cix. fig. 12, it agrees in some respects, but the nucule of 
this species is much more oval and the wing narrow. This specimen may be specifically 
distinct from the other specimens figured by Lesquereux under the same name on 
pi. ex. figs. 6-9. 

Samaropsis quadriovata may be further compared with the Cardiocarpon 
Oliveiranum White, § but one of the distinctive characters of this seed is the notched 
apex of the nucule, which is also slightly narrower in proportion to its length than in 
our seed. With Cardiocarpon Moreiranum White || it has also some similarity, but 
this is a smaller seed, and the nucule appears to be more acute and the wing broader. 

There is no class of fossils more difficult to identify than seeds, for probably their 
form and certainly their size must have varied much at different stages of development, 
and this circumstance is an initial difficulty which can only be overcome when a good 
series of specimens are obtainable. Another difficulty, perhaps not less, is the imperfect 
figures and descriptions that have been given of many of the described species. It is 
therefore not without some reluctance that I apply a distinctive name to this seed, 
but, having failed to identify it with any known species, it seemed the only course to 
follow. 

* Cardiocarpum fluitans, Dawson, Quart. Journ. Geol. Soc, vol. xx. p. 165, pi. xii. fig. 74, 1865. 

t Goal Flora, p. 569, pi. lxxxv. figs. 49, 50. 

% Loc. cit., p. 568, pi. lxxxv. figs. 46, 47. 

§ Comrnissao de Estudos das Minas de Carvao de Pedra do Brazil : Final Report, 1908, p. 565, pi. x. fig. 9. 

|| Loc. cit, p. 563, pi. x. fig. 10. 



156 DR ROBERT KIDSTON ON THE 

Horizons and Localities. — Blue Measures, above Brooch Goal : Jubilee Pit, Sand- 
well Park Colliery, West Bromwich : collected by Mr H. W. Hughes. Hamstead 
Colliery, Great Barr, near Birmingham : collected by Mr Henry Insley. 

Samaropsis acuta L. & H., sp. 

1833. Cardiocarpum aculum, L. & H., Fossil Flora, vol. i. pi. lxxvi. 

-Remarks. — The most constant occurrence of Samaropsis acuta L. & H., sp., with 
Eremopteris artemisisefolia Sternb., sp., early gave rise to the suggestion that they 
belonged to that plant. The late Professor Duns, D.D., published a note in the Proc. 
Roy. Soc, Edin., session 1871-72, p. 692, in which he pointed out the frequent associa- 
tion of these seeds with Eremopteris artemisisefolia ; * but up to the present, as far as 
I am aware, no specimens showing their organic connection have yet been discovered. 

Dr Arber figures some seeds under the name of Cardiocarpus acutus L. & H. 
from the Kent Coal Field ; t but if the figure is an accurate representation of the 
fossils, then his seeds must belong to another species. 

There are in my possession some specimens of Samaropsis acuta L. & H, sp., 
which show considerable details of their internal organisation. These I hope to describe 
at an early date. 

Horizon and Locality. — Roof of Stinking Coal: Clattershall Colliery, Brettell Lane. 

Samaropsis sp. 

Remarks. — This species is closely related to the Cardiocarpon elongatum Newberry. \ 
but is slightly smaller. It is 1*10 cm. long and about 0'70 cm. wide; the pointed 
nucule is 0'80 cm. long and 0'60 wide. The wing is only about 1 mm. wide near the 
base, but increases in width upwards and where it extends past the nucule it is 2 "5 mm. 
broad. The extreme apex of the wing seems to be rounded, but is slightly defective, 
so it cannot be determined whether or not it is emarginate. I have only seen two 
specimens, both collected at the same locality by Mr H. W. Hughes. 

Horizon and Locality. — Blue Measures, above Brooch Coal : Jubilee Pit, Sandwell 
Park Colliery, West Bromwich. 

Tripterospermum Brongniart. 

1881. Tripterospermum, Brongt., Reciter, sur les graines fossiles silicifiees, p. 25. 
Description. — Seed with a thick testa and three very prominent wings. When the 
nut is deprived of the testa it generally presents the form of Trigonocarpus. 

* The specimens on which these observations were made were shown to me by the late Dr Duns. All those with 
the associated Samaropsis acuta were the Eremopteris artemisiasfolia, and one of the specimens which he gave me on 
that occasion is in my collection, No. 769. None of his specimens bore any locality, but from an examination of his 
examples I believe they all came from the Northumberland or Durham Coal Field, as the matrix and general appear- 
ance agree with specimens from that locality. 

t Quart. Journ. Geol. Soc, vol. lxv. p. 29, pi. i. fig. 5. (Enlarged 2 times.) 

X Geol. Survey of Ohio, vol. i. part ii., " Palaeontology," section iii. p. 373, pi. xliii. fig. 5, 1873. Lesqx., Coal Flora, 
p. 567, pi lxxxv. fig. 41, 1879. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 157 

Remarks. — Brongniart founded this genus for the reception of certain seeds with 
their structure preserved, but the above external characters given by him for his genus 
so entirely agree with some seeds T have seen from the South Staffordshire Coal Field 
that I have little hesitation in placing them in Tripterospermum Brongt. 

It is quite possible that when the nucules of Tripterospermum are deprived of their 
testa they might be placed in error in the genus Trigonocarpus. Tripterospermum 
seems, however, to be somewhat rare in Britain. 

Tripterospermum ellipticum Kidston, n. sp. 
PI. XIV. figs, la, lb, lc, Id. 

Description. — Seed elliptical, 6 cm. long and 2*50 cm. wide, with three prominent 
wings of an almost equal width of 5 mm. throughout their whole length ; their rounded 
ends project slightly past the apex of the seed, and at the base they become rounded 
and form a small notch ; " nucleus " about 5 cm. long and 1 cm. wide. 

Remarks. — This specimen is preserved in an ironstone nodule. Owing to the 
carbonaceous matter of the wings having caused weak parts in the stone when split, it 
separated into three portions. These are shown in natural size on PI. XIV. at fig. 1, 
a, b, c ; when they are put together, their relationship to each other is shown at 
fig. Id. The seed is well preserved in so far as showing its form and the width of 
the wings. 

The fossil was collected by the late Mr Henry Johnson, F.G.S., who some years 
ago lent it to me for examination. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Tripterospermum Johnsoni Kidston, n. sp. 
PI. XIV. fig. 2. 



»■ 



Description. — Seed narrow cordate, emarginate at apex, about 3*50 cm. long and 
2 cm. wide. Nucule ovate acute, about 1 cm. wide and 1 '50 cm. long. Seed provided 
with three prominent striated wings about 075 cm. wide at centre of seed, but which 
narrow upwards and extend considerably beyond the apex of the nucule ; wing extend- 
ing beyond base and apex of seed ; emarginate. 

Remarks. — This fossil also occurs in an ironstone nodule, of which only one part has 
been preserved. From the manner in which the nucule occupies the highest point of 
the stone, from which the wings dip down on each side, one can easily observe that the 
specimen is a Tripterospermum and not a Samaropsis. In general character it is very 
similar to Tripterospermum ellipticum, but it is distinguished from that species by its 
smaller size, and above all by its greater width and much broader wings. 



158 DR ROBERT KXDSTON ON THE 

T received this specimen from the late Mr Henry Johnson, jun., after whom I have 
pleasure in naming the fossil. It is shown natural size on PL XIV. fig. 2. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. (K/3573.) 

Polypterospermum Brongt. 

1881. Polypterospermum, Brongt., Recherches sur les graines foss. silicifiees, p. 27. 

Description. — Seed ovoid, obtuse at base and pointed at the summit; in transverse 
section hexagonal, with six more prominent wings, one arising from each angle, and six 
short and truncated wings springing from the sides between the angles. 

The specimens placed under this genus have twelve wings, but in the case of casts 
it may be difficult to determine whether the wings are of equal or unequal width. The 
fact of the seeds possessing twelve wings would seem to be sufficient justification for 
their being included in Polypterospermum, even if the wings were of equal width. 

Polypterospermum ornatum Kidston, n. sp. 
PL X. figs. 6, 7, and 7a ; PL XIV. figs. 5-9. 

Description. — Seed oval, hexagonal in transverse section, from 1*40 cm. to 2 cm. 
long, and bearing twelve wings. The wings are of unequal width, those springing from 
the angles wider than those which arise between the angles. At the apex the wings — 
possibly only the wider ones which spring from the angles — extend upwards and terminate 
in sharp points. Outer surface between the wings ornamented by somewhat irregular 
transverse ridges. 

Remarks. — In a compressed condition it might be very difficult to recognise this 
seed, as in such a state all the wings except those at the margins might be obliterated ; 
but all the specimens described here are preserved in clayband ironstone nodules, and 
though some are compressed, the majority have left an uncompressed impression in the 
nodule, which, when split open, shows the impression of the uncompressed seed. In all 
these cases the seed has been entirely removed, and only its hollow impression remains. 
When the nodules are split, the wing is generally found on the matrix forming a 
border to the two margins of the seed, and the positions of the other wings are indicated 
by longitudinal furrows on the impression. Although, through imperfections, no one 
specimen has shown the twelve wings as far as I know, still, from an examination 
of such specimens as we possess, it would appear that twelve wings was the number 
possessed by the seed. 

Fig. 9, PL XIV., shows what appears to be half of a seed ; the other half of the nodule 
has unfortunately not been preserved. It is slightly compressed, but the position of 
four or five wings can be seen, in addition to one on the matrix at each side of the seed, 
so that twelve seems to have been the original number of wings here. The same seems 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 159 

probable in regard to the seeds seen on PL X. fig. 6 and fig. 7, though there is only 
about the third of the circumference of the seed seen at the latter figure, and on 
PL XIV. at figs. 5, 6. 

At fig, 7, PL XIV., a small flattened example is given, but even here the longitudinal 
lines indicating the position of wings can be seen, and a broad wing extends on the 
matrix from each of the margins. This wing is broader than the wing seen on any of 
the other specimens, and can be compared with that shown on fig. 6 of the same plate. 
It is possible, then, that in this species six of the wings were wider than the other six, 
but it should be pointed out that here this apparent difference in width may be due to 
more perfect preservation and not to a true structural difference. At the apex the wings 
seem to have narrowed as they bent over the top of the seed, where they rose upwards 
as sharp points extending past the apex of the seed. When perfect, what now appear as 
spine-like projections may have united amongst themselves to form the micropylar 
tube. They are seen on the crushed specimen given on PL XIV. fig. 8. 

Polypterospermum is described as hexagonal in transverse section, but one would 
think that the six intermediate wings must have had some effect in producing 
intermediate though perhaps weaker angles. 

Between the ribs, the outer surface is ornamented by irregular transverse ridges, 
which are well seen on the specimen given at PL X. fig. 7, and its enlargement, fig. la, 
and on PL XIV. fig. 5, and its enlargement on PL X. fig. 6. This ornamentation 
consists of transverse ridges of unequal distance apart and generally slightly curved. 

Polypterospermum ornatwn seems to be rare in the South Staffordshire Coal Field, 
but more common in the Derbyshire Coal Field, though, as far as I know, at only one 
locality, where a number of specimens have been discovered by Dr L. Moysey, F.G.S., 
to whom I am indebted for the example shown on PL X. fig. 7, and those on PL XIV. 
figs. 5 and 6, which are figured here to illustrate more fully the structure of the seed, 
which may possibly belong to one of the Pteridosperms, though one cannot speak 
with any certainty as to its relationships. 

The South Staffordshire specimens were collected by Mr H. W. Hughes, F.G.S. 

Horizon and Locality. — Ten -foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Carpolithes Schlotheim. 
Oarpolithes ovoideus G. & B., sp. 

1848. Rhabdocarpus ovoideus, Gopp. and Berger, in Berger, Fruct. et semin., p. 22, pi. i. fig. 17. 

1864. ,, „ Gopp., Foss. Flora d, perm. Form., p. 173, pi. xxvii. figs. 9, 10. 

1871. Rhabdocarpus (?) ovoideus, Weiss, Foss. Flora d. jungst. Stk. u. Rothl., p. 206, pi. xvii. fig. 4, 

pi. xviii. figs. 10-14, 18-21. 

1883. Carpolithus ovoideus, Kidston, Trans. Roy. Soc. Edin., vol. xxxiii. p. 404, pi. xxiii. figs. 7, 8. 
1888. ,, „ Kidston, ibid., vol. xxxv. p, 330, plate, fig. 8. 

1892. Cordaicarpus ovoideus, Zeiller, Flore foss. bassin houil. et perm. d. Brieve, p. 92. 



160 DR ROBERT KIDSTON ON THE 

Horizons and Localities. — 

Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr, 

near Birmingham. Collected by Mr Meachem. 
Above Brooch Coal: Tividale. 
Immediately beloiv Thick Coal: Hamstead Colliery, Great Barr, near 

Birmingham. Collected by Mr H. Insley. 

Lagenostoma Williamson. 

Lagenostoma oblonga Kidston, n. sp. 

PI. VII. figs. 1, la, 2, 2a. 

Description. — Seeds borne in pairs at the extremities of short, dichotomously 
divided branchlets, and surrounded by an oblong cupule 1 cm. long and 3'50 mm. wide, 
which splits into six linear segments about 4 mm. long and about 1 mm. wide, that end 
in sub-acute points. Seed small, oval, 2*50 mm. long, and about 1*50 mm. wide. 




Text-fig. 8. — Lagenostoma, oblonga Kidston. Seed enclosed in cupule, enlarged 3 J times. 

Remarks. — A single specimen of this fructification has been found. It is preserved 
in a small ironstone nodule, and the two halves are seen on PI. VII. figs. 1, 2, natural 
size. These are enlarged 2 times at figs, la and 2a. Owing, however, to the 
colour of the matrix, the fossil does not show up so clearly in the photographs as it 
does on the specimen. I therefore add an outline sketch of the cupule on the right 
of fig. 2. Here the form of the segments into which the cupule splits are better seen. 
Their lower portion is linear, but above they contract into a sub-acute point. On all 
the figures indications of the outline of the seed are seen. In one cupule it lies near 
the base ; in the other, higher up (text-fig. 8) — in the latter case evidently displaced 
from its original position. 

The foot stalks are short and stout for the size of the seed. 

Lagenostoma oblonga is closely related to Lagenostoma Sinclairi Kidston, # but 
is distinguished by its larger size, the oblong, not campanulate form of the cupule, 
and the smaller size of its seeds. In Lagenostoma oblonga the oval seed is 2 '50 mm. 
long and 1'50 mm. broad ; in Lagenostoma Sinclairi it is from 4 to 5 '5 mm. long and 
from 1*5 to 3 mm. in breadth. 

* In Arber, Proc. Roy. Soc. London, vol. B, lxxvi. p. 251, pi. ii. figs. 7-11, 1905. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 161 

The specimen was collected by Mr H. W. Hughes: 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Lagenostoma ? urceolaris Kidston, n. sp. 
PI. XVI. figs. 9, 9a, 10, 10a. 

Description. — Seed urceolate, truncate, about 3 cm. long and 2 '50 cm. wide. 
Micropyle base surrounded by a prominent canopy about 50 cm. high and about 
0*50 cm. wide, and of about the same height as micropyle. 

Remarks. — The seed occurs in an ironstone nodule, and though few of the 
structural details are preserved, those which are shown are of sufficient interest 
for description. 

The two halves of the specimen are given natural size on PI. XVI. figs. 9, 10, and 
enlargements are added at figs. 9a, 10a. The general form of the seed is urn-shaped, 
with a truncate top. The integument is seen on fig. 9a at a, which gradually expands 
upwards to form a prominent "canopy" b. The pollen chamber is seen at c, which 
contracts into the micropylar beak d. All trace of a "central column'' springing 
from the base of the pollen chamber has entirely disappeared, if ever such a structure 
existed in this seed. 

At e are preserved the crumpled and broken-up remains of the nucellus, and at/" 
the empty passage of the vascular strand which entered the chalaza is seen. Between 
the remains of the nucellus and the integument is a space, now filled with white 
calcite g, which looks as if the nucellus might possibly have been free down to the base, 
but probably this space has been formed through the decay of the tissue at this part. 
If, however, the nucellus were free to the base, then of course the seed cannot be 
included in the genus Lagenostoma, where it is provisionally placed. Another 
explanation may be suggested for the parts lettered 6, figs. 9, 10. These may be fleshy 
expansions of a soft outer layer, and not a cupular structure, but this explanation seems 
very improbable. It is much larger than any other member of the genus as at present 
known, but that circumstance is not sufficient for its exclusion. 

Although we may not be able to allocate the seed definitely to any genus, there 
does not seem to be room for much doubt that it belongs to one of the Pteridosperms. 

The specimen was collected by Mr H. W. Hughes, F.G.S. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

t 

Rhabdocarpus Goppert and Berger. 
Rhabdocarpus elongatus Kidston. 

1886. Rhabdocarpus elongatus, Kidston, Trans. Geol. Soc. Glasgow, vol. viii. p. 70, pi. iii. fig. 6. 
1910. „ „ Arber, Proc. Forks. Geol. Soc, vol. xvii. p. 155, pi. xviii. fig. &. 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 21 



162 DR ROBERT KIDSTON ON THE 

Remarks. — Some very fine specimens of this seed have been given me by 
Mr H. W. Hughes, F.G.S., from the Clayscroft Openwork, the largest attaining 
a length of 29 mm. ; but in no other respect than in size do they differ from the type 
specimens, which are only about 15 mm. long. These two extremes in length are, 
however, connected by a chain of intermediate sizes. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

Rhabdocarpus Renaulti Kidston. 

1890. Rhabdocarpus ovoideus, Renault (non Gopp. and Berger), Flore foss. terr. houil. de Commentry, 
p. 639, pi. lxxii. fig. 20. 

Description. — Seed oval, about 4 cm. long and 2 cm. broad, contracted at apex 
into a micropylar beak, and at base into a stalk-like prolongation. Outer surface 
irregularly striated with numerous fibrous thread - like strands, which are very 
irregularly distanced from each other and flexuous in their upward course. 

Remarks. — A single specimen of this seed has been found by Mr H. W. Hughes 
in the South Staffordshire Coal Field, but I possess the same fossil from the Westphalian 
Series of Yorkshire, which was collected by Mr Hemingway (No. 3289). 

As the name given by Renault to his seed had been previously employed by 
Goppert and Berger, # and notwithstanding that the Rhabdocarpus ovoideus Gopp. 
and Berger is now placed in the genus Carpolithes or Cordaicarpus, it is necessary 
to apply another specific name to Renault's Rhabdocarpus, and I propose to dis- 
tinguish it as Rhabdocarpus Renaulti. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

Rhabdocarpus Oliveri Kidston, n. sp. 
PL XVI. figs. 6, 6a, 7, and 8. 

Description. — Seed radiospermic, ovate acute, 4 cm. long (probably longer when 
micropyle was perfect) and 2*10 cm. wide. Outer surface with numerous longitudinal 
striae, at micropylar end gently ribbed and formed of an outer sarcotesta or soft layer, 
and an inner sclerotesta or stony layer, within which is the nucellus. Pollen chamber 
large, broad, and contracted into a micropylar tube. 

Remarks. — Five specimens of this seed have been found at Clayscroft Openwork, 
Coseley, all of which occur in ironstone nodules. Of these, both halves of three of the 
nodules have been preserved. 

On PI. XVJ. figs. 6 and 7 the two halves of the same specimen are shown natural size. 
That at fig. 6 shows some indications of the internal organisation of the seed, which is 

* Fruct. et semin., p. 22, pi. i. fig. 17, 1848. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 163 

held in position by carbonate of lime ; that at fig. 7 shows the impression of the outer 
surface of the seed. In this half there were also originally a few remains of the 
internal structure, but so imperfect that they threw no light on the organisation of the 
seed. They were therefore removed with acid, so that the appearance of the outer 
surface of the seed could be seen. A plastocene cast was made from this, and from it 
the photograph given at fig. 7 was taken. Fig. 8 is from the cast of another specimen, 
from which the contained matter was similarly removed with acid. 

The outer surface of the seed is seen to have numerous longitudinal narrow bands 
which project on the surface ; they are not equally distant from each other, and 
frequently bend somewhat to one side or the other. 

They sometimes run in pairs, but are more commonly single, and converge towards 
the base of the seed. In one case at least these bands seem to unite. On the half 
shown at fig. 8 these bands are seen more distinctly on the cast than in the photo- 
graphs, but are fairly distinctly represented on the figures, especially on fig. 8, where 
there are about thirty of them on the exposed surface. 

These bands more probably represent sclerenchymatous fibres than a vascular 
system, but one cannot speak with certainty on this point. 

On the upper part of the seed, that occupied by the pollen chamber and micropyle, 
the outer surface seems to have been slightly ribbed. 

What remains of the internal organisation can be best understood by referring to 
fig. 6a, which shows the specimen given at fig. 6, enlarged 2 times. 

The micropyle is seen at a, springing from the roof of the pollen chamber b. At c 
is preserved what is probably the upper part of the nucellus. The stony layer or 
sclerotesta is seen at d, at e the remains of the contracted prothallus, and at f the 
cavity originally occupied by the sarcotesta or soft outer layer, now filled with lime. 

The teeth-like points seen on the outer wall of the pollen chamber between the 
letters a and b are sections of the ribs of the sclerotesta on the upper part of the seed, 
broken through obliquely. 

It is impossible to refer this seed, with our present scanty knowledge, to any definite 
plant, but the external characters do not seem to differ in any way from the seeds 
of Neuropteris. They possess the same striated outer surface, and differ only from 
those of Neuropteris heterophylla and Neuropteris obliqua in form and in being 
broader, or more oval. It is slightly smaller than the largest known seed of 
Neuropteris heterophylla, which is about 5 cm. long, while those of Neuropteris 
obliqua are even larger, though their full length is not known. # 

That it is Pteridospermous there seems little reason to doubt. 

I have pleasure in naming this fossil after Professor F. W. Oliver, F.R.S., to whom 
we owe so much for our knowledge of the Carboniferous seeds. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. Collected by Mr H. W. Hughes. 

* See Kidston and Jongmans, Archives Ne'erlandaises d, Sc. exacles et nat., ser. iii., B, vol. i. p. 25, plate, 1911. 



164 DR ROBERT KIDSTON ON THE 

Rhabdocarpus inflatus Lesqx. 

1884. Rhabdocarpus inflatus, Lesqx., Coal Flora, vol. iii. p. 815, pi. ex. fig. 36. 
1890. Carpolithus inflatus, Kidston, Trans. York. Nat. Union, part xiv. p. 63. 

Remarks.— The longitudinal striations are very fine and regular, and in some cases 
are scarcely visible. They appear to be more distinct when the epidermal layer is 
removed from the specimen, and are possibly fine strands of sclerotic tissue. Though 
I was originally inclined to include these fossils in Carpoliihes, the presence of the 
longitudinal bands must exclude them from that genus, as far as at present I 
understand it. 

On account of our imperfect knowledge of the seeds that occur as incrustations, any 
proposed classification can only be regarded as provisional, though it is necessary to 
distinguish the various groups by different provisional generic names, if for no other 
reason than to enable one to record and to refer to them. 

Horizons and Localities. — 

Roof of Brooch Coal : Tividale. 

Roof of Thick Coal : Bradley Colliery, Bilston. 

Rhabdocarpus Wildi Kidston, sp. 

1892. Carpolithes Wildi, Kidston, Trans. Manchester Geol. Soc, part xiii. p. 408 (text-fig.). 

Remarks. — As this species has also the longitudinal parallel bands of Rhabdocajpus, 
I have removed it from Carpolithes, in which genus I originally placed it. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Trigonocarpus Brongniart. 

Trigonocarpus Noeggerathi Sternb., sp. 

1826. Palmacites Noeggerathi, Sternb., Essai flore monde prim., vol. i. fasc. iv. p. xxxv and p. 49, pi. lv. 

figs. 6, 7. 
1828. Trigonocarpum Noeggerathi, Brongt., Prodrome, p. 137. 

1886. Trigonocarpus Noeggerathi, Zeiller, Flore foss. bassin houil. d. Valen., p. 649, pi xciv. figs. 8-11. 

{Excl. ref. L. & H.) 

1888. ,, ,, Kidston, Trans. Roy. Soc. Edin., vol. xxxiii. p. 403, pi. xxiii. 

fig. 3. 

1889. „ ,, Kidston, ibid., vol. xxxv. p. 414, pi. ii. fig. 4. 

1826. Palmacites dubius, Sternb., Essai flore monde prim., vol. i. fasc. iv. p. xxxv and p. 50, pi. lviii. 
figs. 3a, b, c, d. 

Remarks. — This seed is apparently rare in the coal field. 

Horizon and Locality. — Ten-foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 165 

Trigonocarpus sp. 

Remarks. — Several specimens of Trigonocarpus have occurred at the following 
localities, but were not in a good state of preservation for a satisfactory determination, 
though specifically distinct from that mentioned above. 
Horizons and Localities. — 

Ten-foot Ironstone Measures : Ettingshall. 
Immediately below Bottom Coal : Ruiton, near Sedgley. 

Hexagonocarpus Renault. 

1890. Hexagonocarpus, Renault, Flore foss. ten: houil. de Gommentry, deux, part, p. 649. 

Description. — Seeds elongated, hexagonal in transverse section, and prolonged at 
the angles into six more or less prominent wings. 

Remarks. — These seeds are most frequently represented by casts of the cavity 
originally surrounded by the sclerotesta, and which in section show a corresponding 
number of ridges to that of the wings. 

Hexagonocarpus is distinguished from Trigonocarpus in having six equally 
prominent ribs on the " nut " and in the base of the seed ending in a hexagonal flattened 
cone caused by the six ribs extending over its surface and meeting in the centre. 

In Trigonocarpus there are three prominent and three faint ridges, the latter often 
scarcely visible, and the nut has more a circular than a hexagonal form in transverse 
section. The ribs also do not extend over the base, which is generally bluntly rounded. 

Well-preserved specimens of Hexagonocarpus are very easily distinguished from 
Trigonocarpus, especially when the nut is free from the matrix or the base exhibited ; 
but, if badly preserved and partially embedded in the matrix, they might easily be 
mistaken for Trigonocarpus. 

The genus Hexagonocarpus does not appear to be common in British Carboniferous 
rocks, though more than one species is known to occur. 

Hexagonocarpus Hookeri Kidston, n. sp. 

1848. Hooker, "On the Structure and Affinities of Lepidostrobus," Mem. Geol. Survey of Great Britain, 
vol. ii. part ii. p. 456, fig. 5. 

Description. — Nucule 6*50 mm. to 8 mm. long, gradually tapering to a pointed 
apex, and terminating below in a short hexagonal cone. The sides have six equally 
prominent ribs, which extend over the base and divide it into six triangular areas. 

Remarks. — The specimen figured by Hooker as a probable sporangium of 
Lepidodendron elegans is evidently a small Hexagonocarpus. I have seen other 
and better examples of " nuts " which are apparently referable to the same seed, from 
Dove Cliff, near Barnsley, Yorkshire, collected by Mr W. Hemingway (Hor. Woolley 
Edge Rock, Westphalian Series). These latter examples are free from the matrix, 



166 DR ROBERT KTDSTON ON THE 

and show very clearly the distinctive characters between Hexagonocarpus and 
Trigonocarpus. 

There is no definite locality given for the specimen figured by Hooker, but it was 
said to be in the collection of the late Mr John Gray, of Dudley, and occurred in a 
nodule of iron pyrites. 

Hexagonocarpus Hookeri is very rare in the South Staffordshire Coal Field, and 
the only locality from which I have seen specimens is that mentioned below. 

Horizon and Locality. — Ten-foot Ironstone Measures : Clayscroft Openwork, 
Coseley, near Dudley. 

Gymnospermese. 
Whittleseya Newberry. 

The genus Whittleseya has been referred by some to the Ginkgoacese or 
Salisburiacese, and the evidence for and against this position being accorded it has 
been fully discussed by Seward and Gowan, # and by White, t 

It might, however, be stated that this suggested affinity of Wliittleseya rests entirely 
upon the form and structure of the leaves, and therefore does not stand on a sure basis. 

If I am correct in referring the fossils described here under the name of Whittleseya 
l.fevtilis to that genus, then its affinities appear to be rather with the Cycads than 
with Ginkgoacese. 

Whittleseya elegans Newberry. 

1853. Whittleseya elegans, Newberry, Ann. of Sci. of Clevel., vol. v. No. 1, p. 116, figs. 1, 26. (Fide 

Lesquereux.) 
1879. ,, „ Lesquereux, Coal Flora, vol. ii. p. 523, pi. iv. figs. 1, la. 



1885. 
1900. 
1901. 
1904. 
1912. 



Renault, Cours de botan. foss., vol. iv. pp. 69 and 193, pi. v. figs. 9, 10. 
Zeiller, Elements de paleobot., p. 250, fig. 176. 
White, Ottawa Naturalist, vol. xv. No. 4, p. 110, pi. vii. fig. 5. 
Potonie\ Abhild. u. Beschreib. foss. Pflanzen, Lief. ii. No. 40, figs. 1-4. 
Thomas, Palaeobot. Zeitschrift, vol. i. p. 46, text-figs. 1, 2. 



Remarks. — The only British specimen yet discovered, as far as I am aware, is 
that figured and described by Mr H. Hamshaw Thomas, J and which was found by 
Mr S. P. Heath. 

Horizon and Locality. — Ten-foot Ironstone Measures. Contained in an ironstone 
nodule from Doulton's Clay Pit, near Dudley. 

Whittleseya (?) fertilis Kidston, n. sp. 

PL XV. figs. 1 to 10. 

Description. — Sporangial structure formed of two dentate, cuneate, scale-like parts, 
with many strong parallel veins and upper angles slightly rounded and infolded, basal 

* Ann. of Hot., vol. xiv. p. 138 ; see also pp. 135-147. 

t Ottawa Naturalist, July 1901, pp. 106-110 ; see also Scott, Studies, 2nd ed., part ii. p. 611, and Zeiller, Elements 
d. pale'obot., p. 247. J Loc. cit. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 167 

extremity contracted into a pedicel-like termination. In one of the scales is a narrow 
depression becoming more pronounced towards the upper end, in the other a 
corresponding elevation, but not of so great extent as the depression ; thus when the 
two scales rest on each other at their margins a space is left between them, and though 
they meet together closely they do not appear to be united ; the upper toothed margins 
of the two scales also bend inwards towards each other, but also appear to remain free. 
The two scales thus form a closed, somewhat flattened tubular sporangial-like structure. 
Within this sporangial body numerous spores are produced, which, in the fossil state, 
adhere to the inner surface of the two scales. In all cases the sporangia have 
disappeared from the specimens examined. The microspores are from m 210 to m 222 
long, oval, smooth, with thick walls, and sometimes show indications of a central 
longitudinal cleft or line running down one surface of the spore. 

Remarks. — Several specimens of these interesting fossils are figured on PI. XV. to 
show their varying form and size. Their length varies from 1"40 cm. to about 2 "40 cm. 
The width also varies in proportion to the length of the scale ; hence some specimens 
appear much narrower than others. This is seen if figs. 2 and 6 be compared. Their 
upper margin bears six to eight teeth. These bend inwards along with a small part 
of the upper end of the scale to meet the corresponding teeth on the other scale ; hence 
the teeth are not well seen in the photographs, where they are represented in a 
foreshortened view, but they can be observed on figs. 1 to 4 and the corresponding 
enlargements of these specimens. As mentioned already, the tip of the whole scale 
bends inwards as well as the margins ; so when the two scales are in close contact at 
their upper and lateral margins, which they certainly were, though probably not 
organically united to each other, a little pocket is formed between them. The fact 
that the upper margin of the scales had distinct teeth would perhaps indicate that 
this part, though closely adpressed to the same part of its opposite neighbour, 
remained free, and probably any union that took place between the scales was only at 
the base. 

When the small nodules containing these fossils are split — and as far as I know they 
have hitherto only been found in such at Clayscroft Openwork — many are empty, but 
a fair proportion contain microspores. These sometimes lie loose as a powder, but 
when most plentiful they are cemented or held in position on the bract, or perhaps 
more correctly, the sporangiophylls, by carbonate of lime, and are seen to adhere 
equally to the inner surface of both the scales forming the fertile structure. This, 
however, does not necessarily prove that they were originally borne on both surfaces, 
for the lime in solution, while filling the cavity, would distribute any loose spores 
throughout its substance ; and when the nodules are split, the lime adhering to both 
the inner surfaces shows each to be coated with microspores. 

It has already been mentioned that one of the two scales forming the sporangial 
structure has a deep pocket-like depression in the upper part, and it is, I believe, in 
this inner and deeper cavity that the microspores are developed, though the pocket 



168 DR ROBERT KIDSTON ON THE 

practically extends over the whole surface of the scale, though shallow and not so 
pronounced in the lower half. 

A scale entirely covered with microspores embedded in lime is seen natural size at 
fig. 5, and enlarged about 4 times at fig. 5a. Here, as in all the other examples 
seen, the sporangia have entirely disappeared, and their original presence is only 
assumed from the occurrence of the microspores. The upper margin is slightly broken, 
but if this scale be compared in its general form and in the probable distribution of 
the microsporangia over the whole of its surface, its general resemblance to the 
pollen -bearing scale of Cycas is very striking ; and although this resemblance is 
only superficial in some respects, it is probably real in others. It is a real resemblance 
in so far as there is a specialised scale, destined for the production of microsporangia, 
but it differs in these being enclosed in a special receptacle formed by a subtending 
scale, which acts as a protection to the sporangia ; but, notwithstanding this additional 
structure, the fossils seem to show Cycadacean affinities, though here the microsporangia 
are apparently borne on the upper surface of the scale, while in the Cycads they are 
borne on the lower surface. 

The bending-in of the upper part of the scale to make a closed organ is well seen 
at fig. 9, but is observable more or less at all the figures, and it is this bending-in 
which causes the apparent rounding of the upper angles of the scales which, if 
straightened out, would have a more cuneate form than they appear to possess in 
the figures. 

The bracts are strongly striated longitudinally with numerous somewhat irregular 
veins. This character is clearly seen at figs. 6, 6a, and more or less distinctly on all 
the other figures. 

The microspores are oval, have thick, smooth walls, and some show clearly the 
presence of a line on one of their surfaces which passes up the centre but which does 
not quite reach the ends. This may indicate the mode in which the spore opened. 
Three spores are seen enlarged 50 times at fig. 10. 

The next question which falls to be considered is the affinities of these micro- 
sporangial organs, and in doing this it is well to keep clearly before us the outstanding 
characters of these scales, which are their cuneate form, dentate upper margin, and 
strong parallel or slightly fiabellate nervation. They have also a dorsal (or ventral) 
ridge — for it is difficult determining which is the dorsal and which the ventral surface 
of the scales, as they all are separate from their parent axis. Now all these characters 
occur in Whittleseya, and though no specimens of leaves of Whittleseya have been found 
associated with the specimens under discussion, a most characteristic leaf of Whittleseya 
elegans Newberry has been found by Mr S. P. Heath at Doulton's Clay Pit, Netherton, 
near Dudley, on the same horizon, and described and figured by Mr H. Hamshaw Thomas,* 
through whose kindness I am enabled to give a figure of the specimen on PL XV. fig. 11. 
It consists of a broadly cuneate leaf, with central ridge and dentate upper margin. As far as 

* Palaeobot. Zeitschrift, vol. i. Heft i. p. 4(i, figs. 1, 2, 1912. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 169 

I can remember, all the specimens I have previously seen and all the descriptions and 
figures of Whittleseya elegans give the leaf as flat, but this one, preserved in an 
ironstone nodule and free from the pressure which usually brings about a flattening 
of the specimen when preserved in shale, shows the ridge quite distinctly, as seen on 
PI. XV. fig. 11. The differences in general structure between the leaf of Whittleseya 
elegans and the fertile scales described here do not, then, seem greater than would be 
expected to occur between the sterile leaf and one modified for the purpose of fructi- 
fication, and it may be remarked that, though these fertile scales were met with in 
tolerable plenty, they occurred only within a limited and comparatively small area 
of the bed. Their horizontal distribution was very restricted. The same circumstances 
were present in regard to the fertile examples of Neuropteris heterophylla Brongt. 
when the seeds were found, and to the microsporangial specimens of Crossotheca 
(Sphenopteris) Honinghausi Brongt., sp., and in each of these cases the fertile 
specimens may have been derived from a small number or even a single individual 
plant of the species. 

If, then, I am correct in referring these fertile scales to Whittleseya, for which 
course there seems to be a fair amount of evidence, the affinities of that genus would 
point to the Cycadacese rather than to Ginkgo, to which there seems to be a too ready 
reference of almost all the Palaeozoic Gymnosperms as ancestral forms. Affinities can 
rarely be safely traced on vegetative conditions of a species. 

All the examples figured here have been collected by Mr H. W. Hughes, F.G.S. 

Horizon and Locality. — Ten -foot Ironstone Measures: Clayscroft Openwork, 
Coseley, near Dudley. 

Dicranophyllum Grand'Eury. 

1874. Dicranophyllum, Grand'Eury, Compt.es rendus Acad. Sc, vol. Ixxx. p. 1021. 
1877. ,, Grand'Eury, Flore carbon, d. dept. de la Loire, p. 272. 

Description. — Stem ligneous and ramifying irregularly and sparsely. Leaves 
linear, decurrent, expanding at the base, dividing dichotomously into two or more 
linear lobes, which terminate in a point more or less sharp. Nerves parallel to the 
margins of the leaves and dividing by dichotomy. Leaves spirally placed, very 
numerous, bases contiguous and attached by a vertically elongated rhomboidal cushion ; 
erect at the extremities of the branches, spreading on the more aged branches, and on 
old stems spring out at right angles or even bent downwards. 

Male flowers consisting of small scaly ovoid cones placed in the axils of the leaves 
and formed of lanceolate, coriaceous scales with a prominent dorsal keel, their 
extremities so prolonged at the summit that the cone appears to end in a hirsute tuft. 
Microsporangia attached to the enlarged summit of the sporangiophore.* 

Seed small, ovoid, apparently attached in two rows, one on each side of the lower 
and undivided portion of the leaf, which above the fertile region dichotomises once. 

* Dicranophyllum robustum, Zeiller, Bull. Sue. ge'ol. de France, 3 e ser., vol. vi. p. 613, pi. x. fig. 3, 1878. 
TRAJSTS. ROY. SOC. EDIN., VOL. I,. PART I. (NO. 5). 22 



170 DK KOBERT K1DSTON ON THE 

Remarks. — Some place Dicranophyllum in the Ginlcgoacese* but Zeiller thinks 
that it might be better to place it in the Conifer -«,t a position which has been given it 
by Renault.^ Until one has more detailed knowledge of the structure of the fructifica- 
tion, it seems impossible to refer the genus with any degree of satisfaction or certainty 
to either of these groups. 

It is to be remembered that, in dealing with these Palaeozoic Gymnosperms, we may 
be dealing with synthetic types which possessed characters that have now split up and 
passed on to different though perhaps allied genera. § 

An excellent synopsis of the literature and species of Dicranophyllum has been 
given by Reniek,|| some of which are, however, only known from very imperfect 
material. 

Dicranophyllum anglicum Kidston, n. sp. 
PL XIV. figs. 3 and 3a. 

Description. — Leaves close, about 3 "50 cm. long, spirally arranged on stem, and 
dichotomising three or four times into slightly spreading narrow, linear, rigid, sharp- 
pointed segments, from 1*25 mm. to 0*50 mm. wide according to position on leaf. 
Undivided basal portion of leaf about 7 mm. long. 

Nervation at base of leaf not clearly shown, but one vein seems to enter it, which 
dichotomises immediately, the two arms taking a submarginal position, one of which 
passes into each of the two arms of the next bifurcation ; these again bifurcate almost 
immediately on entering the segments, assume a marginal position, and pass into the 
next dichotomy. The terminal segments have only a single central vein ; all the others 
show two marginal veins, the result of a basal dichotomy of the single vein which 
enters them. 

Remarks. — The only specimen of Dicranophyllum which I have met with in 
Britain is that shown natural size on PI. XIV. fig. 3, which is contained in an ironstone 
nodule. The stem is badly preserved and does not show the leaf attachments, except 
of those which spring from the sides, whose relative position, however, clearly indicates 
a spiral arrangement on the stem. The specimen is that of a comparatively young 
branch, and the stem at its widest part is only 0*5 cm. broad. 

The leaves are about 3 "50 cm. long and 1*30 cm. wide, and a line drawn round the 
apices of the segments would give a rhomboidal figure. The segments have been rigid, 
and the arms of the dichotomies form an acute angle with a sharp-pointed sinus 
between them. 

At the base of the leaves the nervation is not well seen, but as far as I can make 
out, only one vein appears to enter the leaf, which immediately on entering bifurcates. 

* Potoni6, Lehrbuch, p. 289. t Elements, p. 255. 

I Flore f oss. terr. houil. de Commentry, part ii. p. 626, 1890. 
§ See also Seward and Gowan, Ann. of Bot., vol. xiv. p. 137, 1900. 
|| Ann. Sue. gSl. de Belgi<iue, vol. xxxiv., Memoires p. M., 193, 1907. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 171 

I am not, however, certain of this, though I do not think there are three veins as in 
Dicranophyllum gallicum Grand'Eury.* 

The arms of the first dichotomy show two marginal veins, but these may arise from 
a dichotomy of a single vein at their base, though I cannot trace this clearly ; but in 
the case of the fourth dichotomy of the leaf it is clearly shown that the single vein 
which enters the segments is the result of a dichotomy of the vein at the extreme base 
of the segment from which they have originated, where they show as marginal veins 
before one passes into each of the ultimate segments of the leaf. 

The rhomboidal form and much-divided leaf will easily distinguish Dicranophyllum 
anglicum from the already known species. A leaf enlarged 2 times is given at 
3a. I am indebted to the late Mr D. Rogers for this specimen, from whom I 
received it in 1887, but have delayed describing the plant in the hope that additional 
examples might be discovered, but such, as far as I am aware, has unfortunately not 
been the case. 

Horizon and Locality. — Roof of Brooch Coal: Shut End, near Dudley. 

ROOTLETS. 

Pinnularia Lindley and Hutton. 

Pinnularia columnaris Artis, sp. 

1825. Hydatica columnaris, Artis, Antedil. Phyt., pi. v. 

]886. Pinnularia columnaris, Zeiller, Flore foss. bassin houil. d. Valen., p. 404, pi. lvii. fig. 3. 

1901. ,, ,, Kidston, Proc. YorJcs. Geol. and Polytech. Soc., vol. xiv. part ii. pp. 204 

and 225, pi. xxxv. fig. 1. 
1899. Radicites columnaris, Zeiller, Etude sur la flore foss. d. bassin houil. d'Heraclee, p. 69. 
1848. Asterophyllites Artisi, Gb'pp., in Bronn (pars), Index palxont., p. 122. 
1869. Asterophyllites foliosa, Roehl (non L. & H.), Foss. Flora d. Steinlc.-Form. Westph., p. 24, pi. v. 

fig- I- 

1877. Roots and rootlets, Lebour, Illustr. of Fossil Plants, p. 21, pi. x. 

Horizons and Localities. — 

Blue Measures, six feet above Brooch Coal : Hamstead Colliery, Great Barr, 

near Birmingham. 
Ten-foot Ironstone Measures : Clayscroft Openwork, Coseley, near Dudley. 

Pinnularia capillacea L. & H. 

1834. Pinnularia capillacea, L. & H., Fossil Flora, vol. ii. pi. cxi. 

1858. „ ,, Lesqx., in Rogers, Geol. of Pennsyl., vol. ii. part ii. p. 878 (? pi. xvii. 

fig. 22). 
1869. „ „ Roehl, Foss. Flora d. Steink.-Form. Westph., p. 27 (? pi. ii. fig. 5a), 

pi. iv. fig. 11 (? fig. la). 
1893. Radicites capillacea, Potonie, Flora d. Rothl. v. Thiiringen, p. 261, pi. xxxiv. fig. 2. 
1877. Rootlets, Lebour, Illustr. of Fossil Plants, p. 113, pi. lix. 

* See Zeiller, Ve'get. foss. d. terr. houil, p. 158, 1880. 



172 DR ROBERT KIDSTON ON THE 

Horizons and Localities!. — 

Above Brooch Coal: Grets Green, near West Bromwich. 
Ten-foot Ironstone Measures : Coseley, near Dudley ; Ettingshall. 
Roof of Thick Coal: Bradley Colliery, Bilston. 
Roof of New Mine: Merry hill Colliery, Brierley Hill. 



APPENDIX. 

Annularia microphylla Sauveur. 
PL X. figs. 1-3. 

1848. Annularia microphylla, Sauveur, Veget. foss. d. terr. houil, de la Belgique, pi. Ixix. fig. 6. 

Description. — Stem jointed, finely striated ; internodes on the ultimate branchlets 
short, 4 to 6 mm. long and about 0*75 mm. thick. Leaves as long as the internodes, 
14 to 16 in a whorl, sickle-shaped, lanceolate, widest at their middle, gradually 
narrowing into a sharp point. Midrib very prominent and placed in a distinct 
furrow. Margins of leaf involute. Cones small, terminal, or given off the stem singly, 
apparently not in whorls. (K. No. 2122.) 

Remarks. — Sauveur figured two small fragments of his Annularia microphylla, 
one of which is a terminal portion and the other a piece of a small branch broken over 
at both ends. On both of his figures the leaves are distinctly bent or sickle-shaped, 
and narrower than those of Annularia galioides in proportion to their length. His 
figure, however, gives the idea that the leaves were somewhat blunt-pointed, and it 
was this which made me originally think his figure was a slightly inaccurate drawing 
of Annularia galioides L. & H., sp. 

The lowest whorl of leaves on the left-hand specimen (which is given in inverted 
position) illustrates well the bent and pointed leaves of the plant I now identify as 
Sauvkur's species. The absence of a description to his figures makes the identification 
of some of his plants very difficult. 

Some small specimens of Annularia microphylla Sauveur are given on PL X. 
That seen at fig. 1, natural size, shows the upper part of an ultimate branch. The 
internodes at the lower part are about 6 mm. long, but at the upper end are very much 
shorter. The larger whorls of leaves are 90 mm. in diameter, the individual leaves 
being about 40 mm. long and at their widest part 075 mm. wide. In springing from 
the stem they first assume an almost horizontal direction, but soon gradually bend 
upwards towards the stem. The whorl, as a whole, is thus saucer-shaped. The midrib 
is very prominent and lies in a little furrow, and the margins of the leaves are distinctly 
roiled in (involute) (fig. la). The base of a cone-like structure is attached to the apex 
of this branch, but it is not well preserved. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 173 

Fig. 2 shows portion of three parallel leafy branches which appear to spring from 
the stem lying at their base, but I cannot observe any actual connection. This 
specimen is enlarged 2 times at fig. 2a, where the bent-up leaves with their incurved 
margins, and the prominent midrib placed in a furrow, can all be seen. It is also 
noticeable on this specimen that the leaves decrease in size towards the base of the 
branchlet (fig. 1). 

Another small specimen is given at fig. 3, and enlarged 2 times at fig. 3a. The 
bowl- or saucer-shape form of the leaf whorls is well seen on the enlargement fig. 3a. 

If the two species be now compared, it will be seen that in Annularia microphylla 
Sauveur the leaves are lanceolate and sharp-pointed, with involute margins and a 
prominent midrib placed in a furrow. In Annularia galioides the leaves are much 
broader in proportion to their length and do not end in such prolonged points. The 
midrib is not placed in a furrow and the leaves are flat.* 
Horizons and Localities. — 

Westphalian Series; Barnsley Thick Coal: Monckton Main Colliery, near 

Barnsley, Yorkshire ; and Wolley Colliery, Darton, near Barnsley. 

Collected by Mr W. Hemingway. 

Bensham Seam, Jarrow, County of Durham. Specimen preserved in " Hutton 

Collection," Newcastle-on-Tyne. 

Lanarhian Series : Furnace Bank Pit, Old Sauchie,near Alloa, Clackmannanshire. 

Notes on the Vertical Distribution of the Fossil Plants. 

In all 154 species are recorded from the Westphalian Series of the South 
Staffordshire Coal Field. Of these, 13 are only generically recorded, as the material 
at my disposal was not in a sufficiently good. state of preservation for a satisfactory 
identification or description. 

Twenty new species are described, namely, Sphenopteris deltiformis, Sphenopteris 
Kilimlii, Coseleya glomerata, Pecopteris hepaticwformis, Neuropteris Carpentieri, 
Palseostachya minuta, Sphenophyllum tenuissimum, Sigillaria punctirugosa, Lepido- 
carpon Westphalicum, Samaropsis quadriovata, Tripterospermum ellipticum, Triptero- 
spermum Joh7bsoni,Polypterospermum ornatum, Lagenostoma oblonga, Lagenostoma (?) 
urceolaris, Rhabdocarpus Renaulti, Rhabdocarpus Oliveri, Hexagonocarpus Hooheri, 
Whittlesey u (?) fertilis, and Dicranophyllum anglicum ; while the 10 following have 
not previously been recorded for Britain, as far as I am aware : Sphenopteris 
Schatzlariana Stur, sp. (emend.), cf. Sphenopteris Sancta - Felicis Stur, sp., 
Sphenopteris Souichi Zeiller, Crossotheca Crepini Zeiller, Adiantites sessilis Koehl 
[pro. var.), Alethopteris integra Gothan, sp., Ccdamostachys Solmsi Weiss, Huttonia 
spicata Presl, sp., Lycopodites Meehi Lesqx., and Sigillaria cordigera Zeiller. 

Although a considerable number of species have been met with in the South 

* Note. — For references to Annularia galioides L. & H., sp., see ante, p. 121. 



174 DR ROBERT KIDSTON ON THE 

Staffordshire Coal Field, many of them appear to be very rare, though perhaps they 
may be much more common than is at present suspected, for from many localities 
I have few or no records, and from some of the coal seams, especially in the northern 
area of the coal field, there are no records at all. 

The publication of this paper may call the attention of some to the deficiency of 
records from many of the coal seams, and I would ask such to lose no opportunity 
for collecting specimens, so that we may gain a more accurate conception of the 
vertical and horizontal distribution of the fossil plants of this coal field. 

The following table contains a list of al] the species found in the South Staffordshire 
Coal Field, and gives their vertical distribution. The list is very typical of the 
Westphalian Series. 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 



175 



Bottom Coal. 






x • ■ 


Intermediate Measures. 




X 








Fireclay Coal. 




X 


x • • 


New Mine Coal. 


x x X 


X 




Stinking Coal = 
Sulphur Coal. 




White Ironstone. 




Rubble Coal. 




Heathen Coal. 




Gubbin's Ironstone. 




e3 
<D 
H 

<j 

a 

<v 
+3 
o 

0) 

o 
O 
k<] 

H 

C4-I 
O 

s 

"3 


Bottom Benches Coal. 




Slipper Coal. 




Sawyer's Coal. 




PatchelPs Coal. 




Stone Coal. 




John Coal. 






Foot Coal. 




Brassil's Coal. 




Benches Coal. 




Tow Coal. 




Lamb's Coal. 




Jay's Coal. 




Top Slipper Coal. 






Thick Coal. 


XX -X -XXX • -XX • • -X • 


x ; 


< XX- 


Brooch Coal. 






X • X 


Little Two-foot Coal. 




Upper Sulphur Coal. 





s s 



,0 a> 



o 



pq . 

1? e 



<d ~<S> 12 
8 "!» ■•» 



5- ■*- 



a • 



T3 *S 
^ o 






a 

re; 



OQ ^ 






w 3 

S-S a 

a ."> *■» 

2 a H 
a <u S 

'&."£« 



t«V2 X 






«2 






cu 



be 

a 
o 












l.§ 



03 oq 



s 
a 
a? 



^•SOj^ 



be o " 
c -^ 

f-H T3 

"<s> ■ — i 

s S 3 
|>8 a 

" -3 nS "«* 

a «s» Bh 
£ >2 ^ 






o 



176 



DR ROBERT KTDSTON ON THE 



S 

o 
ID 



so 
• <^ 

"fe. 

«o 
•*» 

CO 

g 



Bottom Coal. 


• ■ ■ • • • • • X ■ • X X X • ■ • 


Intermediate Measures. x • • x x x • • • 


Fireclay Coal. 


x • • • x ■ ■ • • x ■ • x x .... 


New Mine Coal. 


x x • • X X X • ■ • 


Stinking Coal = 
Sulphur Coal. 




White Ironstone. 


x 


Rubble Coal. 


I 


Heathen Coal. 




Gubbin's Ironstone. 


aj 

< 

a 

CD 

f4 

O 

_d 

15 < 
o 
O 

M 

H 

o 

"S 

'3 


' Bottom Benches Coal. 




Slipper Coal. 




Sawyer's Coal. 




Patchell's Coal. 




Stone Coal. 




John Coal. 




Foot Coal. 




Brassil's Coal. 


Benches Coal. 




Tow Coal. 




Lamb's Coal. 




Jay's Coal. 




Top Slipper Coal. 


Roofs Coal. 




Thick Coal. 


X X • • • X • X X XXX X X xxxx 


Brooch Coal. • xxx • • x • • x • • x x xx-x 


Little Two-foot Coal. 




Upper Sulphur Coal. 






Sphyropteris. 

obliqua Marrat, sp. 
Zeilleria. 

Avoldensis Stur, sp. 

delicatula Sternb., sp. 

re. sp. .... 
Eremopteris. 

artemisixfolia Sternb., sp. . 
Adiantites. 

sessilis Roehl., pro. var. 
Archseopteris. 

Reussii Ett., sp. . 
Telangium. 

asteroides Lesqx., sp. 
Qoseleya. 

glomerata Kidston, n. sp. . 
Pecopteris, 

Miltoni Artis, sp. . 

Volkmanni Sauveur 

hepaticseformis Kidston, n. sp. 
Dactylotheca. 

pluinosa Artis, sp. . 
Mariopteris. 

muricata Schl., sp. 
Alethopteris. 

lonchitica Schl., sp. 

decurrens Artis, sp. 

Davreuxi Brongt., sp. 

valida Boulay 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 177 



x • x x • x ■ • x x • • x • x 



«5 



lO 

cq X-X-X 



N - - x ' * X • X 



CO 



<N X • X 



<M 



O 
CM 



OS 



CD 

lO 

1— I 



i -# X X • X X X-XXXXXX-X- XX X X X-X-XXX XXXX XX 

_ _ 

eoxx- X • XXX-XX-X-X •• • • XXX--X-- X--- x • 

<M • • • • • • • • • .... . . 



a, 



05 






?!!i'|Hi!ti|Hlfl|.vi,i .liliifi I .Hflt-.il 



SOS; ^l^KjO ^ ^ 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 23 



CSS 



178 



DR ROBERT KIDSTON ON THE 



^3 

HO 

o 



CO 

CO 

■fe. 

CO 
CO 

CO 





















Intermediate Measures. 


• • • -X • • • X • X • • 


■- ■- • • 


X 


Fireclay Coal. 


X • X X • • 


X • 




New Mine Coal. 


• - * X • • XX-- • X • • 


X X X • • • 




Stinking Coal = 
Sulphur Coal. 














White Ironstone. 














Rubble Coal. 














Heathen Coal. 




Gubbin's Ironstone. 


X • X X • • 










CO 
< 

a 

CO 

XI 
■*^ 

o 

00 

a 

-2 < 
o 
O 

co 

H 

*4-l 

o 

a 

CO 

'3 

a 1 










Slipper Coal. 














Sawyer's Coal. 














Patchell's Coal. 














Stone Coal. 














John Coal. 














Foot Coal. 














Brassil's Coal. 














Benches Coal. 















Tow Coal. 














Lamb's Coal. 














Jay's Coal. 














Top Slipper Coal. 






















U"™ 1 " wwna. 








Thick Coal. 


X • • X • . e- . . . XX- 


X -X • ■ °* 


X 


Brooch Coal. 


•x x-x 


< X ■ X X • • 


X 


Little Two-foot Coal. 

































u 

CO 

M 

eo 

N 



M 



J) IB g 



CO 

a 



T3 



X> 



w 





sphenophyl 
stellata Scl 
mostaehys. 
Solmsi We 






ya. 
gshau 
ata Pi 


g Ph 




ta Ste 

lum. 

foliun 




a, a. 


Ettin 
elong 




■«a 


s^ a, s 


s 

B 






Ca 




© 


8 


« 










4o 




a 


3 






Oh 




A 


£ 



CT 1 



a a 






X 






h9 § 

-w « S S s 

a,^ fe 'e a,-s 



CO 



S © 



o ® 

CO M 

-» » 
CC S 

gl 

i-J 

§13 






CO 
ON) 



CN 



o 
o> 

I-H 

co 

r-l 

t~ 

I— I 

CO 

T— I 

lO 

I— I 

rH 

CO 

I— I 



OS 
CO 

CO 

CO 
CM 






-c 



•^ 



a. 






FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 



179 



X • X x x • x • • x x 


x X • • • 


XX X • XX-- 


•• • X • X --XX-XX-X-'-X. X ■ • 


x • 


x x 




































•X • • • X ••••XXX X • •• X XX-- X 


XX x • x X X ■ • ■ X X • ■ X- X •• • -XX- X 






Geinitzi Schimper . 

triangularis Zeiller 
Lepidophyllum. 

intermedium L. & H. 
Lepidophloios. 

laricinus Sternb. . 
Halonia. 

tortuosa L. & H. 
Bothrodendron. 

minutifolium Boulay, sp. . 
Sigillaria. 

discophora Kbnig., sp. 

semipulvinata Kidston 

elegans Brongt. 

cf. nodosa Bowman, sp. 

trigona Sternb., sp. 

tessellata Brongt. . 

mamillaris Brongt. 

scutellata Brongt. . 

vulgaris Artis, sp. . 

cordigera Zeiller 

■reniformis Brongt. . 

elongata Brongt. 

rugosa Brongt. 

punctirugosa Kidston, n. sp. 
Sigillariostrobus. 

ciliatus Kidston 

sp. .... 
Gyperites. 

bicurinata L. & H. 
Stigmaria. 

flcoides Sternb., sp.f 

reticulata Gopp., pro. var. . 
Lepidocarpon. 

Westphalicum Kidston, n. sp. 
Cordaites. 

borassifolius Sternb., sp. 

principalis Germar, sp. 

palmsefornis Gopp., sp. 

Brandlingi AVitham, sp. . 
Artisia. 

approximata Brongt., sp. . 






to 









CO 






O 



to 



^3 



O 



as 

CO 
tO 

m 

CO 



180 



DR ROBERT KIDSTON ON THE 





Bottom Coal. 






Intermediate Measures. 






Fireclay Coal. 






New Mine Coal. 






Stinking Coal = 

O .... ...V. .. • * >. 

Sulphur Coal. 




White Ironstone. 






Rubble Coal. 






Heathen Coal. 






Gubbin's Ironstone. 






<D 
f-. 

<J 

a 

u 
o 

CD 

r\ 

1 ' 
O 

M 
o 

13 

H 

o 

a 


1 Bottom Benches Coal. 






Slipper Coal. 






Sawyer's Coal. 






Patchell's Coal. 






Stone Coal. 




"6 

3 


John Coal. 




•<s> 

o 


Foot Coal. 




Brassil's Coal. 




=o 


Benches Coal. 




CO 


Tow Coal. 






Lamb's Coal. 




^ 


Jay's Coal. 




•<s> 

CO 


Top Slipper Coal. 




^ 

g 


Roofs Coal. 




CO 


Thick Coal. 


XX- X---- XX X X XX XXXXX X 




Brooch Coal. 


• x • x xxx-x •• • x X • 




Little Two-foot Coal. 






Upper Sulphur Coal. 








' ' A ' ' 'ft 

Oh • . bD &. 
o" £3 OT ^ CL, ft • M Oh Ph 

* w ' • Jj|i" gg a 1 tg ga«^ 1 

«,j :s3 w b 3| jI a sa 3-31 g.f a 

|5 ' ' c?"|« •g'JfliB 3 3,* AHsjl | 

111 -iilli Ifl ilji I fi 111 III II 

i i 1 # | i 1 I 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 



181 



cS x 



<M - - - - ■ • 

<M ..... 

« x 

CO 

<M 

Cq • . • . . . . 

r— I 

c<, • . . . . . 

o 

(M • • • • • • ■ 

Oi 

T— I * " " " * * 

00 

I— I " " * • 

to 

w 

i— i 

CO 

i— i ..... 

i— I ..... 

i—l 

O ... 

Oi ■ ■ ■ ■ . . 
CO 

t- . . . . . . . 

<o . . . . . . 

>o 



"* X X XX • XX 
CO . . • • X X X 
<M 



s-i 

o 



T3 



M 






32 ?3 






«3 ^ 



g 



« e o 
*o w ^ 
s 
a 

8 



d cq 



182 



DR ROBERT KIDSTON ON THE 









INDEX. 








PAGE 


PAGE 


Adiantites . . . . . . . 95 


Carpolithes ....... 159 


sessilis Roehl, pro. var. . 






95 


ovoideus Gbpp. & Berger 






159 


Alethopteris .... 






101 


Gordaianthus 






153 


Davreuxi Brongt., sp. . 






102 


Pitcairnise L. & H., sp. . 






153 


decurrens Artis, sp. 






102 


Volkmanni Ett., sp. 






153 


Grandini Brongt., sp. . 






103 


sp. 






153 


integra Gothan, sp. 






103 


sp. 






153 


lonchitica Schl., sp. 






101 


Gordaites .... 






150 


Serli Brongt., sp. . 






103 


borassifolius Sternb., sp. 






150 


valida Boulay 






102 


Brandlingi Witham, sp. 






152 


Annul aria .... 






121 


palmx/ormis Gbpp., sp. 






151 


galioides L. & H., sp. . 






121 


principalis Germar, sp. . 






151 


microphylla Sauveur 






172 


Coseleya .... 






97 


radiata Brongt. 






121 


glomerata Kidston 






97 


sphenophylloides Zenker, sp. 






122 


Grossotheca .... 






90 


stellata Schl., sp. . 






123 


Crepini Zeiller 






91 


Aphlebia .... 






115 


Honinghausi Brongt. 






90 


crispa Gutbier, sp. 






115 


Hughesiana Kidston 






91 


sp. 






116 


Gyperites .... 






146 


Archseopteris 






95 


bicarinata L. & H. 






146 


Reussii Ett., sp. . 






95 


Dactylotheca 






99 


Artisia .... 






152 


plumosa Artis, sp. 






99 


approximate), Brongt., sp. 






152 


Dicranophyllum . 






169 


Asterophyllites 






120 


anglicum Kidston . 






170 


charxformis Sternb., sp. 






121 


Equisetites .... 






117 


equisetiformis Schl., sp. . 






120 


Heminginayi Kidston . 






117 


grandis Sternb., sp. 






120 


Eremopteris 






95 


longifolius Sternb., sp. . 






120 


artemisixfolia Sternb., sp. 






95 


Bothrodendron 






138 


Halonia .... 






137 


minuti/olium Boulay, sp. 






. 138 


tortuosa L. & H. . 






137 


Boweria .... 






89 


Hexagonocarpus . 






165 


Schatzlarensis Kidston . 






90 


Hookeri Kidston . 






165 


Catamites .... 






117 


Huttonia .... 






128 


Britannicus Weiss 






119 


spicata Sternb. 






128 


Cisti Brongt. 






118 


Lagenostoma 






160 


ramosus Artis 






119 


oblonga Kidston . 






160 


Schutzei Stur 






118 


urceolaris Kidston 






161 


Suckoivi Brongt. . 






117 


Lepidocarpon 






147 


undulatus Sternb. . 






118 


Wcstphalicum Kidston . 






148 


Waldenburyensis Kidston 






118 


Lepidodendron 






132 


sp. (tuber) . 






119 


aculeatum Sternb. 




134 


Calamostachys 






123 


acutum Sternb. 




135 


Solmsi Weiss 






123 


cf. distant Lesqx. . 




135 


sp. ..... 






125 


obovatum Sternb. . 


134 


sp. 






125 


ophiurus Brongt. . 






132 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 



183 











PAGE 










PAGE 


Lepidodendron. 


Samaropsis ....... 154 


simile Kidston . . 134 


acuta L. & H., sp. 






. 156 


Lepidophloios 






137 


Gutbieri Geinitz, sp. 








154 


laricinus Sternb. . 








137 


Meachemi Kidston 








154 


Lepidophyllum 








137 


quadriovata Kidston 








155 


intermedium L. & H. 








137 


sp. 








156 


Lepidostrobus 








136 


Sigillaria .... 








138 


Geinitzi Schiiuper 








136 


cordigera Zeiller . 








144 


triangularis Zeiller 








136 


discoplwra Kdnig., sp. 








138 


variabilis L. & H. 








136 


elegans Brongt. 








139 


Lonchopteris 








105 


elongata Brongt. . 








145 


rugosa . 








105 


mamillaris Brongt. 








li3 


Lycopodites . 








132 


cf. nodosa Bowman, sp. 








139 


Meeki . 








132 


punctirugosa Kidston 








145 


Mariopteris . 








100 


reniformis Brongt. 








145 


muricata Schl., sp. 








100 


rugosa Brongt. 








145 


Neuropteris . 








106 


scutellata Brongt. . 








143 


Carpentieri Kidston 








112 


semipidvinata Kidston 








138 


gigantea Sternb. . 








108 


tessellata Brongt. . 








142 


Grangeri Brongt. . 








111 


trigona Sternb., sp. 








140 


heterophylla Brongt. 








106 


vulgaris Artis, sp. 








143 


obligua Brongt., sp. 








111 


Siyillariostrobus . 








146 


Osmundx Artis, sp 








112 


ciliatus Kidston . 








146 


rarinervis Bunbury 








110 


sp. 








146 


Scheuchzeri Hoffm. 








112 


Sphenophyllum 








129 


Schlehani Stur 








111 


cuneifolium Sternb., sp. 








129 


tenuifolia Schl., sp. 








108 


tenuissimum Kidston 








129 


sp. 








115 


sp. 








131 


Odontopteris 








105 


Sphenopteris 








78 


alpina Sternb., sp. 








105 


artemisixfolioides Crepii 


l 






83 


Paleeostachya 








125 


deltiformis Kidston 








85 


elongata Presl 








127 


dilatata L. & H. . 








80 


Ettingshauseni Kidston 








125 


furcata Brongt- 








83 


gracillima Weiss . 








127 


Kilimlii Kidston . 








86 


minuta Kidston . 








127 


Laurenti Andrae . 








82 


Pecopteris . 








98 


obtusiloba Brongt. 








78 


hepaticeeformis Kidston 








99 


quadridactylites Gutbiei 








85 


Miltoni Artis, sp. . 








98 


cf. Sancta-Felicis Stur 








84 


Volkmanni Sauveur 








99 


Sauveuri Crepin . 








81 


Pinnularia . 








171 


Schatzlarensis Stur, sp. 








89 


capillacea L. & H. 








171 


Schatzlariana Stur 








84 


columnaris Artis, sp. 








171 


Schillingsi Andrae 








82 


Polypterospermum 








158 


Souichi Zeiller 








88 


ornatum Kidston . 








158 


spinosa Gopp. 








83 


Renaultia . 








89 


trifoliolata Artis, sp. 








81 


gracilis Brongt., sp. 








89 


Walteri Stur, sp. . 








88 


Khabdocarpus 








161 


Sphyropteris 








91 


elongatus Kidston . 








161 


obliqua Marrat, sp. 








91 


inflatus Lesqx., sp. 








164 


Spiropteris . 








116 


Oliveri Kidston 








162 


sp. 








116 


Renaulti Kidston . 








162 


Stigmaria . 








147 


Wildi Kidston 








164 


ficoides Sternb., sp. 








147 



184 



DR ROBERT KIDSTON ON THE 



PAGE 



PAGE 



Stigmaria. 




Tripterospermum. 








reticulata Gopp. .... 


147 


Johnsoni Kidston . . .157 


Telangium ....... 


96 


Whittleseya . 






166 


asteroides Lesqx., sp. . 


96 


elegans Newberry . 




. 


166 


Trigonocarpus ..... 


164 


fertilis Kidston 






166 


Noeggerathi Sternb., sp. 


164 


Zeilleria 






92 


sp. 


165 


Avoldensis Stur, sp. 






92 


Tripterospermum ..... 


156 


delicatula Sternb., sp. 






94 


ellipticum Kidston 


157 


sp. 






94 



EXPLANATION OF PLATES. 

Plate V. 
Fig. 1. Sphenopteris dilatata L. & H. 

Fragment of pinna. Natural size. Locality — Clayscroft Openwork, Coseley, near Dudley. 
Horizon — Ten-foot Ironstone Measures. Specimen in the Collection of British Museum, 
Geological Department. (No. V. 1377.) 
Fig. la. The same specimen, enlarged 2 times. 
Fig. 2. Sphenopteris Schillingsi Andrae. 

Termination of pinna. Natural size. Same locality and horizon as last. 
Fig. 3. Sphenopteris Kilimlii Kidston. 

Fragment of a frond. Natural size. Locality — Doulton's Clay Pit, Netherton. Horizon — Blue 
Measures, six feet above Fireclay Coal. (K. No. 1613.) 
Fig. 4. Coseleya glomerata Kidston. 

Small fragment, showing the sporangia clustered round the rachis. Locality — Clayscroft 
Openwork, Coseley, near Dudley. Horizon — Ten-foot Ironstone Measures. 
Fig. 4a. Coseleya glomerata Kidston. 

Same specimen as the last, enlarged 2 times. At the broken pinna marked a the sporangia 
are seen surrounding the rachis on all sides. 
Fig. 4&. Coseleya glomerata Kidston. 

Some sporangia from the part lettered a in fig. 4a, enlarged about 9 times to show form of 
sporangia. 
Fig. 5. Coseleya glomerata Kidston. 

A small fragment, enlarged 2 times to show the form of the sporangia. 
Fig. 6. Coseleya glomerata Kidston. 

Fragment of a pinna, enlarged 2 times to show spiral or verticillate arrangement of sporangia 
at a. Same locality and horizon as last. 
Fig. 7. Archxopteris Reussii Ett., sp. 

Fragment of frond. Natural size. Locality — Tividale, Dudley. Horizon — Above Brooch Coal. 
Specimen in the Johnson Collection, Geological Department, British Museum. (No. V. 
1366.) 
Fig. 7a. Archxopteris Reussii Ett., sp. 

Some pinnules, enlarged 2 times to show their nervation. 



Plate VI. 

Fig. 1. Neuropteris gigantea Sternb. 

Upper portion of pinna, showing terminal pinnules. Natural size. Locality — Clayscroft Open- 
work, Coseley. (K. No. 4016.) 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 185 

Fig. 2. Neuropteris gigantea Sternb. 

Single pinnule, enlarged 1\ times to show nervation. Same locality and horizon. 
Fig. 3. Neuropteris gigantea Sternb. 

Upper part of a pinna, showing the two terminal pinnules. Natural size. Same locality and 
horizon. (K. No. 4014.) 
Fig. 4. Neuropteris gigantea Sternb. 

Specimen showing a (?) double dichotomy of the pinna rachis. Natural size. Same locality and 
horizon. (K. No. 207.) 
Fig. 5. Neuropteris gigantea Sternb. 

Small portion of a partially developed frond. At a the main rachis is densely covered with 
scales. Natural size. Same locality and horizon. (K. No; 3760.) 
Fig. 6. Neuropteris gigantea Sternb. 

Portion of a circinately coiled pinna, shown natural size. The rachis is densely covered with 
scales. Same locality and horizon. (K. No. 3761.) 
Figs, la and 7b. Neuropteris gigantea Sternb. 

The two halves of the same nodule, natural size, showing a young pinna circinately coiled. 
The rachis is very densely covered with scales. Same locality and horizon. (K. Nos. 4117 
and 4118.) 
Fig. 7c. Neuropteris gigantea Sternb. 

The specimen shown at 7b, enlarged 2 times. 
Fig. 8. Spiropteris sp. 

Young frond, showing circinate vernation and dichotomy of rachis. Natural size. Same locality 
and horizon. (K. No. 3334.) 
Fig. 9. Spiropteris sp. 

Very young, complete frond, showing thick petiole with scale scars, bifurcating at the apex, 
the two arms being circinately coiled. Natural size. Same locality and horizon. 
(K. No. 3321.) 
Fig. 9a. Spiropteris sp. 

Apex of last specimen, enlarged 2 times. 

Specimen from which fig. 4 is taken was received from the late Mr Henry Johnson, that repre- 
sented at fig. 9 from Mr W. Madblby, and all the others on this plate from Mr H. W. 
Hughes, F.G.S. 



Plate VII. 

Figs. 1 and 2. Lagenostoma oblonga Kidston. 

Two seeds enclosed in their cupules. Natural size. The seeds are seen at a and b. Locality — 
Clayscroft Openwork, Coseley. Horizon — Ten-foot Ironstone Measures. 
Figs, la and 2a. Lagenostoma oblonga Kidston. 

The same specimen, enlarged 2 times. 
Figs, 3 and 4. Neuropteris heterophylla Brongt. 

Fully developed and young seed. Natural size. Same horizon and locality. 
Fig. 5. Zeilleria Avoldensis Stur, sp. 

Seed-bearing frond. Natural size. Same locality and horizon. 
Fig. 5a. Zeilleria Avoldensis Stur, sp. 

Portion of last, enlarged 2 times to show the spread-out cupules and the central mamillae to 
which the seeds were attached. 5b. The cupule seen at a, fig. 5a, enlarged 6 times. 
Fig. 6. Zeilleria Avoldensis Stur, sp. 

Portion of microsporangia-bearing frond. Natural size. Locality — Tividale. Horizon— Roof 
of Brooch Coal. 
Fig. 6a. Zeilleria Avoldensis Stur, sp. 

Portion of same specimen, enlarged 2 times. 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 5). 24 



186 DR ROBERT KIDSTON ON THE 

Fig. 7. Pecopteris ? hepaticatformis Kidston. 

Small portion of a pinna. Natural size. Locality — Coseley 1 Horizon — Ten-foot Ironstone 
Measures. 
Fig. 7a. Pecopteris ? hepaticxformis Kidston. 

A few pinnules, enlarged 10 times. 

Plate VIII. 
Neuropteris Carpentieri Kidston. 
Locality — Claysoroft Openwork, Coseley. Horizon — Ten-foot Ironstone Measures. 

Figs. 1, 2, 4, and 7. Natural size. 

Figs. 3a, 3b, 4a, 5a, 5b, 6, and 7a, enlarged 2 times. 

Fig. 8. Portions of three "columns" of microspores, showing part of the complete spore contents of a 

sporangium. x 50. 
Fig. 8a. Microspores, one showing triradiate ridge. x 500. 
For explanation of other figures, see text. 

Plate IX. 

Fig. 1. Annularia stellata Schl., sp. 

Portion of branch. Natural size. Locality — Jubilee Pit, Sandwell Park Colliery, West 
Bromwich. Horizon — Blue Measures, six feet above Brooch Coal. 
Fig. la. Annularia stellata Schl., sp. 

Portion of specimen enlarged 2 times. 
Fig. 2. Catamites tuber. 

Natural size. Locality — Clayscroft Openwork, Coseley. Horizon — Ten-foot Ironstone 
Measures. 
Fig. 3. Palxostachya Ettingshauseni Kidston. 

Cone. Natural size. Locality — Mount Pleasant, Brierley Hill. Horizon — Roof of New 
Mine Coal. 
Fig. 3a. Palxostachya Ettingshauseni Kidston. 

Part of cone, enlarged 2 times. 
Fig. 3b. Palxostachya Ettingshauseni Kidston. 

Microspores enlarged 50 times. Fig. 3c. Microspore enlarged 500 times. 
Fig. 4. Galamostachys Solmsi Weiss. 

Part of cone attached to its pedicel. Natural size. Locality — Clattershall Colliery, Brettell 
Lane. Horizon — Roof of New Mine Coal. 
Fig. 4a. Calamostachys Solmsi Weiss. 

Part of cone, enlarged 2 times to show form of bracts. 
Fig. ib. Calamostachys Solmsi Weiss. 

Microspores eidarged 50 times. Fig. 4c. Microspore enlarged 500 times. Fig. 4o\ Megaspore 
enlarged 50 times. 

Plate X. 
Fig. 1. Annularia microphylla Sauveur. 

Showing branch terminating in a cone. Locality — Furnace Bank Pit, Old Sauchie, near Alloa, 
Clackmannanshire. Horizon — Lanarkian Series. Natural size. (K. No. 92.) 
Fig. la. Annularia microphylla Sauveur. 

Portion of same specimen, enlarged 2 times to show form of leaves. 
Fig. 2. Annularia microphylla Sauveur. 

Small specimen, shown natural size. Locality — Monckton Main Colliery, near Barnsley, York- 
shire. Horizon — Thick Coal, Westphalian Series. (K. No. 2123.) 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 187 

Fig. 2a. Annularia microphylla Sauveur. 

Same specimen as last, enlarged 2 times to show form of leaf and nervation. 
Fig. 3. Annularia microphylla Sauveur. 

Fragment of branch. Natural size. Same locality and horizon as at fig. 2. 
Fig. 3a. Annularia microphylla Sauveur. 

Portion of same specimen, enlarged to show the central vein. 
Fig. 4. Coseleya glomerata Kidston. 

Small specimen, enlarged 2 times to show bramble-like appearance of masses of sporangia. 
Locality — Clayscroft Openwork, Coseley. Horizon — Ten-foot Ironstone Measures. 
Fig. 5. Sphenophyllum sp. 

Group of four united sporangia. Natural size. 5a. The same specimen, enlarged 2 times. 
Locality — Jubilee Pit, Sandwell Park, West Bromwich. Horizon — Blue Measures, six feet 

above Brooch Co;d. 
Fig. 6. Polypterospermum ornatum Kidston. 

Portion of seed, enlarged '2 times to show ornamentation. Locality — Shipley Clay Pit, Ilkeston, 
Derbyshire. Horizon — Below Top Hard Coal. Westphalian Series. (K. No. 4651.) 
Fig. 7. Polypterospermum ornatum Kiilston. 

Part of seed. Natural size. Same horizon and locality. (K. No. 4650.) 
Fig. la. Polypterospermum ornatum Kidston. 

Same specimen, enlarged 2 times to show the ornamentation. 
Fig. 8. Cf. Sphenopteris Sanda-Felicis Stnr, sp. 

Small fragment of a pinna. Natural size. Locality — Doulton's Clay Pit, Netherton. Horizon — 
Roof of Fireclay Coal. 
Fig. 8a. Cf. Sphenopteris Sanda-Felicis Stur, sp. 

Same specimen, enlarged 2 times to show pinnule segmentation. 
Fig. 9. Sphenopteris deltiformis Kidston. 

Fragment of frond. Natural size. Locality — Race Course Pit, Round Oak. Horizon — 
Whitestone. 
Fig. 9a. Sphenopteris deltiformis Kidston. 

Some pinnse from same specimen, enlarged 2 times to show form and segmentation of pinnules. 

Plate XI. 
Fig. 1. Palxostachya minuta Kidston. 

Small slab showing many specimens scattered over its surface. Natural size. Locality — 
Doulton's Clay Pit, Netherton. Horizon — Between Fireclay Coal and Bottom Coal. 
(K. No. 4476.) 
Fig. la-lc. Palxostachya minuta Kidston. 

Small fragments of cones enlarged 2 times, from the same specimen. 
Fig. 2. Cone of Lepidodemlron ophiuras Brongt. 

Specimen showing cone attached to leafy branch. Natural size. Locality — Clayscroft Open- 
work, Coseley. Horizon — Ten-foot Ironstone Measures. (K. No. 912.) 
Fig. 3. Cone of Lepidodendron ophiurus Brongt. 

Another specimen longitudinally broken through the centre and filled in with carbonate of lime, 
showing the axis and bracts. Natural size. Same locality and horizon. (K. No. 3765.) 
Fig. 4. Huttonia spicata Sternb. 

Portion of a cone. Natural size. Locality — Doulton's Clay Pit, Netherton. Horizon — Blue 
Measures, six feet above Fireclay Coal. 

Plate XII. 
Fig. 1. Sigillaria, cf. nodosa Bowman, sp. 

Fragment of cortex, showing outer surface. Natural size. Locality — Merryhill Colliery, Brierley 
Hill. Horizon — Roof of New Mine Coal. Lighting parallel with ribs. (K. No. 3562.) 



188 DR ROBERT KIDSTON ON THE 

Fig. la. Sigillaria, cf. nodosa Bowman, sp. 

Portion of same specimen, enlarged 2 times, with illumination at right angles to ribs, to show 
the form of the leaf scar. 
Fig. 2. Sigillaria trigona Sternb., sp. 

Natural size. Locality — Tipton. Horizon — Roof of Thick Coal. 
Fig. 2a. Sigillaria trigona Sternb., sp. 

Portion of same specimen, enlarged 2 times to show form of cushion and leaf scar. 
Figs. 3 and 4. Sigillaria trigona Sternb., sp. 

Two outline drawings of cushion and leaf scar to show opening of ligule pit and diverging lines 
on base of cushion. 
Fig. 5. Sigillaria cordigera Zeiller. 

Natural size. Locality — Mount Pleasant, Brierley Hill. Horizon — Roof of New Mine Coal. 
Fig. 5a. Sigillaria cordigera Zeiller. 

From same specimen, enlarged 2 times to show the leaf scars and surface ornamentation of 
cortex. 
Fig. 6. Sigillaria punctirugosa Kidston. 

Specimen, natural size. Locality — Mount Pleasant, Brierley Hill. Horizon — Roof of New 
Mine Coal. (K. No. 3474.) 
Fig. 6a. Sigillaria punctirugosa Kidston. 

A small portion, enlarged 2 times to show form of leaf scar and ornamentation of ribs. 



Plate XIII. 

Figs. 1-6. Lepidocarpon Westphalicum Kidston. 

Locality — Clayscroft Openwork, Coseley. Horizon — Ten- foot Ironstone Measures. 



Figs. I and 2. Natural size. 
Fig. 3. Enlarged 2 times. 
Fig. 4. Enlarged 4 times. 



Fig. 5. Enlarged 4 times. 
Fig. 6. Enlarged 8 times. 



For description, see text. 



Plate XIV. 



Figs, la, lb, and \c. Tripterospermum ellipticum Kidston. 

The seed seen on three of the surfaces into which the nodule split, and which corresponds to the 
three pieces of the nodule lettered a, b, c, fig. Id. Natural size. Locality — Clayscroft 
Openwork, Coseley. Horizon — Ten-foot Ironstone Measures. 
Fig. 2. Tripterospermum Johnsoni Kidston. 

Portion of seed, showing two of the wings. Natural size. Same locality aud horizon as last. 
(K. No. 3573.) 
Fig. 3. Dicranophyllum anglicum Kidston. 

Portion of small branch with leaves attached. Natural size. Locality — Shut End, near 
Dudley. Horizon — Roof of Brooch Coal. Collected by Mr D. Rodgebs. (K. No. 4623.) 
Fig. 3a. Dicranophyllum anglicum Kidston. 

Leaf from same specimen, enlarged 2 times. 
Fig. 4. Huttonia spicata Sternb. 

Portion of specimen given on PI. XL fig. 4, enlarged 2 times to show bracts. 
Fig. 5. Polypterospermum ornatum Kidston. 

Portion of seed seen natural size, showing ornamentation and longitudinal furrows indicating 
position of wings. Locality — Shipley Clay Pit, Ilkeston, Derbyshire. Horizon — Below 
Top Hard Coal. (K. No. 4651.) 



FOSSIL FLORA OF THE STAFFORDSHIRE COAL FIELDS. 189 

Fig. 6. Polypterospermum ornatum Kidston. 

Part of another specimen showing ornamentation on surface between the wings and the narrow 
wing on the matrix at the right-hand side. Natural size. Same locality and horizon 
as last. (K. No. 4648.) 
Fig. 7. Polypterospermum ornatum Kidston. 

Small specimen with wings. Natural size. Locality — Clayscroft Openwork, Coseley. 
Horizon — Ten-foot Ironstone Measures. 
Fig. 8. Polypterospermum ornatum Kidston. 

Crushed specimen showing upward prolongation of the wings beyond the apex of the seed. 
Natural size. Same locality and horizon as last. 
Fig. 9. Polypterospermum ornatum Kidston. 

Large crushed specimen showing the furrows originally occupied by the wings, and their points 
projecting past the apex of the seed. Natural size. Same locality and horizon. 



Plate XV. 
Figs. 1-10. Whittlesey a ? fertilis Kidston. 



Fig. 5a. Enlarged 4 times. 

Fig. 6. Natural size. 

Figs. 6a and 6/;. Enlarged 2 times. 

Figs. 7, 8, and 9. Enlarged 2 times. 

Fig. 10. Microspores enlarged 50 times. 



Fig. 1. Natural size. 

Figs, la and lb. Enlarged 2 times. 

Fig. 2. Natural size. 

Figs. 2a and 2b. Enlarged 2 times. 

Figs. 3 and 4. Natural size. 

Fig. 5. Natural size. 

For description, see text. 
Fig. 11. Whittlesey a elegans Newberry. 

Leaf natural size. Locality — Doulton's Clay Pit, Netherton. Lent by Mr H. Hamstead 
Thomas. 



Plate XVI. 

Fig. 1. Samaropsis quadriovata Kidston. 

Seed natural size. Locality — Jubilee Pit, Sandwell Park, West Bromwich. Horizon — Blue 
Measures, six feet above Brooch Coal. (K. No. 3770.) 
Fig. la. Samaropsis quadriovata Kidston. 

The same seed, enlarged 2 times. 
Fig. 2. Samaropsis Gutbieri Geinitz, sp. 

Natural size. Locality — Clayscroft Openwork, Coseley. Horizon — Ten-foot Ironstone Measures. 
Figs. 3 and 4. Sphenopliyllum tenuissimum Kidston. 

Two halves of nodule. Natural size. Locality — Clayscroft Openwork, Coseley. Horizon — 

Ten-foot Ironstone Measures. 

Fig. 4a. Sphenophyllum tenuissimum Kidston. 

Same specimen, enlarged 2 times. 

Fig. 5a. Sphenophyllum tenuissimum Kidston. 

Group of united sporangia, of which three are shown but the fourth is broken off. Enlarged. 
From same specimen. 
Fig. 6. Rhabdocarpus Oliveri Kidston. 

Natural size. Locality — Clayscroft Openwork, Coseley. Horizon — Ten-foot Ironstone Measures. 
Fig. 6a. Rhabdocarpus Oliveri Kidston. 

The same specimen, enlarged 2 times, a, micropyle ; b, pollen chamber ; c, upper part of 
nucellus ; d, stony layer of seed (Sclerotesta) ; e, remains of contorted prothallus ; /, cavity 
left through decay of soft layer of seed (Sarcotesta), now filled with lime. 
TRANS. ROY. SOG. EDIN., VOL. L. PART I. (NO. 5). 25 



190 DR KIDSTON ON THE FOSSIL FLORA OF STAFFORDSHIRE COAL FIELDS. 

Fig. 7. Rhabdocarpus Oliveri Kidston. 

Other half of the same specimen after removal of the lime and other matter adhering to the 
matrix, as seen from a photograph of a plastocene cast of the impression in the matrix 
showing striated onter surface. Natural size- 
Fig. 8. Rhabdocarpus, Oliveri Kidston. 

Photograph of a plastocene cast of the impression of another specimen to show the longitudinal 
striations. Natural size. Same locality and horizon. 
Figs. 9 and 10. Lagenostoma? urctolaris Kidston. 

The two halves of the same specimen, shown natural size. Locality — Clayscroft Openwork, 
Coseley. Horizon — Ten-foot Ironstone Measures. 
Figs. 9a and 10a. Lagenostoma 2 urceolaris Kidston. 

Same specimen, enlarged 2 times, a, integument; b, canopy; c, pollen chamber; cl, micropyle, 
e, remains of nucellus ; /, passage through which the vascular bundle enters the seed ; 
g, 1 space between cupule and seed. 



Trans. Roy. Soc. Edin>. Vol. L.— Plate V. 

Kidston: Fossil Flora of the Staffordshire Coal Fields — Part III. 




R. Kidston, Photo. 



46 -f 



M'Farlane & Erskine, Lith., Eilin. 



1. Sphenopteris dilatata, L. & H. 2. Sphenopteris Schillingsi, Andrse. 3. Sphenopteris Kilimlii, Kidston. n. sp. 
4-6. Coseleya gloinerata, Kidston. n. sp. 7. Archseopteris Reussii, Ettingshausen sp. 



ijins. Roy. Soc. Edin^. Vol. L.— Plate VI. 

Kidston : Fossil Flora of the Staffordshire Coal Fields — Part III. 




Photo 



M'Farlane & Erskine. Lith., Ellin 



1-7. Neuropteris gigantea, Sternb 8-9 Spiropteris. 



rans. Eoy. Soc. Edin r . Vol. L. — Plate VII. 

Kidston : Fossil Flora of the Staffordshire Coal Fields — Part III. 




ba -f 



E. Kidston, Photo. 



M'h'ailane 4 Erskine, Litta., Edin. 



1-2. Lagenostoma oblonga, Kidston. n.sp. 3-4. Neuropteris heterophylla, Brongt. 
5-6. Zeilleria Avoldensis, Stur. sp. 7. Pecopteris hepaticseformis, Kidston. n. sp. 



Trans. Roy. Soc. Edin-. Vol. L.— Plate VIII. 

Kidston : Fossil Flora of the Staffordshire Coal Fields— Part III. 




la -f 



R. Kidston, Photo. 



M'Farlanc & Erskine, Lith., Edin. 



Neuropteris Carpentieri, Kidston. n.sp. 



>ans. Roy. Soc. Edim. Vol. L. — Plate IX. 

Kidston : Fossil Flora of the Staffordshire Coal Fields — Part III. 




3a -f- 



ia 



M'Farlaue & Erskine, Lith., Edin. 



Ustou, Photo. 



1. Annularia stellata s Schl. sp. 2. Calamites tuber. 3. Palaeostachya Ettingshauseni, Kidston. 



Vans. Roy. Soc. Edim Vol. L. — Plate X. 

Kidston : Fossil Flora of the Staffordshire Coal Fields — Part III. 




8a -f- 



MF.irlane & Erskiue. Lith., EiliD 



1-3. Annularia microphylla, Sauveur. 4. Coseleya glomerata, Kidston.n.sp. 5. Sphenophyllum sp. 
6-7. Polypterospermum ornatum, Kidston. n. sp. 8. Sphenopteris cf . Sancti-Felicis, Stur sp. 



m 



m 



"rans. Roy. Soc. Edin*. Vol. L. — Plate XL 

Kidston : Fossil Flora of the Staffordshire Coal Fields — Part III. 




II idston. Photo. 



MFarlaue k Erskine, Lith., Ed in. 



_ 



1. Palaeostachya minuta, Kidston. n . 8 p. 2-3. Lepidodendron ophiurus, Brongt. 4. Huttonia spicata, Sternb. 



>ans. Roy. Soc. EdiiK Vol. L.— Plate XII. 

Kidston : Fossil Flora of the Staffordshire Coal Fields — Part III. 








s 


1-1 ' * '^^ImlHSBl 

if j't'SiiSHI' hH 
















B 

5): 




■k ' ^fe J< 






tjidston, Photo. 



MFarlane & Erskine Lith.. Edin. 



1. Sigillaria cf. nodosa, Bowman, sp. 2-4. Sigillaria trigona, Sternb. sp. 
5. Sigillaria cordigera, Zeiller. 6. Sigillaria punctirugosa, Kidston. n. 8P . 



r ra „, Roy. Soc. Editf Vo] l _ Plate xnI 

Kidston : Fossil Flora of the Staffordshire Coal Fields— Part III. 







s 






sp 



sp 






3 -f 



M'Farlane 4 Erekine, Lith., Edin. 



Lepidocarpon Westphalicum, Kidston. u, , 



Trans. Roy. Soc. EdhV. Vol. L. -Plate XIV. 

Ktdston : Fossil Flora of the Staffordshire Coal Fields— Part III. 




M'Farlane & Erskine, Lith., Edin. 



1. Tripterospermum ellipticum, Kidston. n.sp. 2. Tripterospermum Johnsoni, Kidston. n . «.. 
DicranophjUum anglicum, Kidston. „. , P . 4. Huttonia spicata, Sternb. 5-9. Polypterospermum ornatum, Kidston. n. 8J 



>ans. Roy. Soc. Edin r . Vol. L.— Plate XV, 

Kidston: Fossil Flora of the Staffordshire Coal Fields— Part III. 




2« -f 



M'Farlane k Erskine. Lith.. Edin 



1-10. Whittleseya (?) fertilis, Kidston. u. sp . 11. Whittleseya elegans, Newberry 



W Roy. Soe. Edin'. Vo , ^^ ^ 

Kidston : Fossil Flora of the Staffordshire Coal Fields— Part III. 




•i ton, Photo. 



M'Farlane & Erskine, Lith., Edin. 

Sa^ropsis quadriovata, Kidston, „. 2. Sama ropsis Gutbieri, Geinitz 8 p. 3-5. SphenophyUun, tenuissinnnn, Kidston. .. ,, 
6-8. Rhabdocarpus Oliveri, Kidston.n* P 9-10. Lagenostoma 1 urceolaris, Kidston. »..p. 



( 191 ) 



VI. —The Anatomy of a New Species of Bathydoris, and the Affinities of the 
Genus : Scottish National Antarctic Expedition. By T. J. Evans, M.A. (Oxon.), 
Lecturer in Zoology in the University of Sheffield. Communicated by Dr J. H. 
Ash worth. 

(MS. received October 3, 1913. Read January 19, 1914. Issued separately April 1, 1914.) 

[Plates XVII. and XVIIL] 

Introductory. 

The genus Bathydoris was created by Bergh in 1884 in his Eeport on the 
Nudibranch Mollusca collected by the Challenger. In his account of the anatomy 
of the new genus Bergh draws attention to the anomalous combination of characters 
possessed by the animal, and gives it an annectent position between the Dorids and 
the Tritonids, but places it among the Dorids on account of the predominance of Dorid 
features. The single specimen of Bathydoris abyssorum was dredged off New South 
Wales in 2425 fathoms. A second specimen of this peculiar genus was obtained by 
the Danish Ingolf Expedition and described by Bergh in 1900. This specimen came 
from 1870 fathoms in Davis Strait, and resembled B. abyssorum, with specific 
variations. Thus Bathydoris came to be regarded as an isolated genus with the 
characters of a connecting link, and appropriately a denizen of deep water. Our 
anatomical knowledge of the animal is derived almost entirely from Bergh's accounts 
of the two species mentioned, and is moderately extensive, considering the rather 
imperfect state of preservation of the material and the fact that he was dependent on 
single specimens in each case. 

Two more specimens were brought from the Antarctic by the Discovery and 
described by Sir Charles Eliot in the Report of the National Antarctic Expedition 
published by the Natural History Museum in 1907. These constituted separate 
species resembling Bergh's, but their state of preservation was such that Eliot was 
able to add little to our knowledge of the anatomy of Bathydoris. The discovery of 
one of Eliot's species in 100 fathoms dispels the idea that the genus is confined to 
the great depths. 

A specifically distinct specimen was also taken by the German Antarctic Expedi- 
tion, but Thiele, in the Report on the Mollusca recently published, confines himself 
to a superficial description in order to retain the rare animal intact as a museum 
specimen. 

It would seem to be the fate of polar expeditions to bring back single specific 
specimens of this genus ; for the species which forms the subject of the present 
memoir is based on one specimen dredged in 1410 fathoms by the Scotia in March 
1904. When the zoological material of the Scottish National Antarctic Expedition 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 6). 26 



192 MR T. J. EVANS ON 

was distributed for identification, the animal went astray, and Sir Charles Eliot, who 
received the Nudibranchs, comments with surprise on the complete absence of Dorids. 

Dr Bruce has provided the following notes on the haul in which the animal was 
included: "Locality, 71° 22' S., 16° 34' W. Bottom temperature, 31 0, 9 F. Surface 
temperature, 29° 9 F. Depth, 1410 fathoms. Bottom, blue mud." The haul included 
specimens of nearly every group of animals from sponges to fishes. 

It is proposed to name the new species Bathydoris brownii, in honour of Dr R. N. 
Rudmose Brown, naturalist to the expedition. To him and to Dr Bruce the author's 
thanks are due for permission to undertake the investigation of its anatomy. 

The six known species of Bathydoris may be tabulated as follows : — 

~ 1. B. abyssorum, Bergh. 2425 fathoms, off New South Wales. ,: Challenger." 

2. B. ingolfiana, Bergh. 1870 fathoms, in Davis Strait. " Ingolf." 

3. B. hodgsoni, Eliot. 100 fathoms, off Coulman Island. "Discovery." 

4. B. inflata, Eliot. Depth not stated, off Coulman Island. "Discovery." 

5. B. clavigera, Thiele. Depth not stated, Gauss Station. " Gauss." 

6. B. brownii, sp. nov. 1410 fathoms, off Coats Land. "Scotia." 

The specimen was preserved in about 5 per cent, formaldehyde, and suffered very 
little distortion or contraction. The viscera were also in excellent condition for 
dissection, and even the histological preservation was found to be remarkably good 
when certain tissues were cut and stained for identification. 



External Features. 

The animal, as preserved, was 75 mm. long, 40 mm. broad, and 35 mm. in height. 
A thin, flabby foot margin extended about 8 mm. beyond the body all round and 
was rather bluntly pointed behind, while anteriorly its thickened edge ran transversely 
across the body behind the head, and had a deep glandular furrow extending into the 
lateral margin for some distance. The contraction of the foot when the animal was 
killed probably accounts for the distension of the dorsal integument and the extruded 
condition of the genital organs, which were forced out to the extent usual among 
Dorids during copulation. The mouth lies entirely on the ventral aspect, and is 
surrounded by crinkled lips surmounted by a bulging forehead, which extends laterally 
into rather long cylindrical oral tentacles. Nearly a third of the length of the ventral 
surface is occupied by the buccal region, the enormous size of which is a striking 
feature of the genus. The arched dorsal surface is covered by an integument which is 
delimited all round the body by a slight fringe at a distance of 4 or 5 mm. from the 
foot. Laterally this fringe or reduced mantle edge is represented by little more than 
a linear thickening of the skin. The dorsal integument is thin, transparent, and 
destitute of spicules, but, as in hodgsoni, injiata, and clavigera, the whole surface seems 
to have been originally studded with papillae. Those that remain on exposed parts 



THE ANATOMY OF A NEW SPECIES OF BATHYDORIS. 193 

of the back are smallest ; while round the dorsal tentacles and along the notaeal 
margin they are much larger. They seem to be more numerous in brownii than in 
any of the other three species possessing them. They are undoubtedly caducous, and, 
as recorded by Eliot, the points of attachment of detached papillae are marked on bare 
places by small circles with a spot in the centre. The only other conspicuous features 
of the dorsal surface are the dorsal tentacles and the anal complex, consisting of the 
anus, the renal pore, and the branchiae. The rhinophores are club-shaped and perfoliated 
all round. There is no trace of a pocket. The branchiae are two in number, and are 
placed symmetrically in front of the prominent anal papilla. Both numerically 
and structurally the branchiae appear to differ very considerably from those of all the 
other species of Bathydoris. Thus, B. abyssorum has five and ingolfiana ten arranged 
in a circle in front of the anus, while Eliot's species have eight and five or six 
respectively similarly arranged. The two tufts in B. brownii are united by a ridge 
which, on dissection, is found to contain the afferent and efferent vessels of the 
branchiae. As the structure of the gills will be described later, it is sufficient here to 
mention that they are not fine dendritic structures like the gill-plumes represented by 
Bergh. The anal papilla is much shorter than in the other species, but this is probably 
due to the relatively greater local distension of the hinder end of the specimen. The 
renal papilla is inconspicuous, and lies in the median line between the anus and the 
base of the gills. 

The extruded genitalia stand out conspicuously on the right side between the 
notaeal margin and the foot, and are thrown further back than in the Dorids by the 
great size of the buccal region. 

Anatomical Description. 

The importance of Bathydoris in auy discussion of the interrelationships and 
classification of the Nudibranchs was recognised by both Bergh and Eliot, and, during 
the examination of the anatomical structure of the present species, this importance 
became increasingly impressive as system after system was considered. This was 
deemed to justify a more complete and detailed account of the anatomy than has 
hitherto been given, especially as facilities for such an account were amply provided in 
virtue of the excellent preservation of the specimen. In the course of this memoir it 
will be noted that the description disagrees with those of Bergh and Eliot on matters 
the bearings of which are of great theoretical interest. These contradictions are some- 
times so striking that the inclusion of the species in the genus Bathydoris seemed 
jeopardised. The points of agreement, however, form such convincing evidence of 
generic identity that the serious divergencies here given must be regarded as corrections, 
based on examination of a more favourable specimen, of observations partially frustrated 
by the poor condition of the material observed by previous workers. 

The various systems, will now be considered in order. 



194 MR T. J. EVANS ON 



The Alimentary System. 



It will be seen in fig. 2 that the general plan of the alimentary system of Bathydoris 
resembles that of the typical Doric!, but a detailed examination shows that there are 
very important differences. The great size of the buccal mass (b.m.) was emphasised 
by Bergh as a characteristic feature of the genus. Indeed, Bergh claimed its supposed 
relationship with the Tritonids solely on the character of the buccal mass and the radula. 
The mouth is surrounded by two sets of lips, an outer and an inner. The outer lips are 
crinkled and fleshy, and leave a wide gape into which the inner lips project. These are 
merely the thickened rim of the outer integument limiting the buccal opening. The 
inner lips {p.m., figs. 2 and 3) are a pair of lateral pads enclosing a relatively small open- 
ing leading into the buccal cavity. The pads have the consistency and appearance 
of hyaline cartilage Histological examination shows that the hyaline substance is a 
cuticular secretion which is continued in varying thickness as a lining of the alimentary 
tube from the buccal rim to the stomach. The underlying epithelium consists through- 
out of tall slender columnar cells, and corresponding columns of the secreted matter are 
faintly visible in the cuticle. 

The buccal cavity contains a pair of dark-brown horny jaws (J., fig. 3) supported on 
muscular pads (P.J., fig. 3) which separate them from the globular odontophore (o.) 
occupying the middle of the floor. The free edges of the jaws are quite blunt, and each 
is produced into a slight beak opposite the mouth ; so that they are probably used as 
a prehensile organ. The mouth must be capable of far more extension than might be 
supposed from its preserved state in order to bring the jaws into action. The radula 
(R.) is narrow in front and broad behind where it enters the radula sac, and the sac is 
entirely contained in the substance of the odontophoral mass. The first row of teeth has 
three teeth on either side of the rhachidian tooth and the radula broadens to 90. 1. 90, 
the formula of the youngest row. The total number of rows is about 50. The rhachidian 
tooth (Rh., fig. 4) has a broad base on which stands a backwardly directed cusp. The 
laterals (L. 1 to L. v ) are formed on the same plan, but the cusp bends from the base 
towards the middle line. The first four laterals differ from the rest in having shorter 
and blunter cusps. Unlike those of the other species, the extreme laterals show little 
sign of reduction. The teeth are firmly fixed in the specially thick cuticular covering 
of the radula mass, which is itself bilobed with a deep median depression. The direction 
of the cusps and of the underlying muscles suggests that the radula is used for 
gripping the food during trituration. The approximation of the cusps of the first 
laterals thus entailed would also explain the broken and irregular appearance of the 
rhachidian cusps as well as the reduction in length of the cusps of the first few laterals. 
Comparison with the figures published by Bergh and Eliot shows that the present 
species can be identified by its radula alone. 

The oesophagus (ce. and cr., fig. 2) seems to differ from that of the other species in 
that it turns to the left, even at its origin from the buccal mass. It is a broad, sigmoid 



THE ANATOMY OF A NEW SPECIES OF BATHYDORIS. 195 

tube with muscular walls lined, as already mentioned, by a thick cuticle. At its lower 
end it ends in a thin-walled sac, the stomach (st., fig. 5), lying below the liver, and 
partially imbedded in its substance. The lining of the oesophagus is, throughout its 
length, thrown into twelve raised longitudinal bands (b.o. , fig. 5) which are covered with 
minute, blunt, brown cones of various sizes. As shown in fig. 6, the cones are partly 
imbedded in the cuticle of which they are specially resistent local modifications. The 
dark tint of the cuticle of the first half of the oesophagus and the extensive crinkling 
of its walls obscured the presence in it of the longitudinal bands which were only seen 
after clearing. Eliot, who describes the denticulate bands in the second half of the 
oesophagus, may have overlooked them in the first half for the same reason. The same 
writer, in his account of B. hodgsoni, names the two regions of the oesophagus the first 
and second stomach — names which seem inappropriate for a thickly cuticularised tube 
which serves merely for the delivery of the food to the sac in which it is actually 
digested. The denticulate cuticle would, moreover, be quite ineffectual for the purpose 
of mastication, and probably serves as a protective layer against the coarse diet of mud, 
sponge, and small shells The thin-walled stomach stands at the junction of oesophagus, 
intestine {int., fig. 5), and main liver ducts (l.t.) into the expanded ends of which the 
food enters for some distance, as in Dorids generally. At its junction with the oeso- 
phagus the stomach has a small pocket (s.r.) like that of the Dorids, which was not seen 
in any of the other species. Its function is not known, though such inapplicable names 
as " pancreas " and " gall-bladder" have been applied to it by different authors. 

It may be mentioned that the similar stomach recess in Doris tuberculata secretes 
a glassy, refringent substance which is also found as a granular deposit on the mucosa 
of the intestine and on the massed Halichondria spicules passing down that tube. This 
suggests a protective function for the organ, its secretion acting as a lubricant for the 
passage of spiculose excrement down the intestine. The liver (I., figs. 2 and 5) is a 
bulky organ which is not invaded by gonad or kidney, and is unlobed except in so far 
as the intestine and the lower end of the oesophagus lie in furrows on its surface. The 
intestine is a rather broad, smooth tube making an arc round the pericardium and end- 
ing by a sphinctered opening on the anal papilla. 

The alimentary tract contained one large piece of undigested sponge, and sponge 
spicules were present in all parts of the stomach and intestine. There were also found 
much mud, bits of old shells, small pebbles, and the spines of Echinids. The animal is 
therefore probably an omnivorous feeder, though the prevalence of sponge suggests that 
it has predilections for that group, like the Dorids. 

The salivary glands {s.g., fig. 2) are fiocculent and voluminous, forming a mass on 
each side pressed against the wall of the oesophagus, and opening by stout ducts on 
the hinder wall of the buccal cavity. Bergh and Eliot mistook them for the blood 
glands, but their histological structure puts their salivary nature beyond doubt. 
Moreover, the true blood glands were found elsewhere. 



196 MR T. J. EVANS ON 

The Nervous System. (Fig. 7.) 

The brain of Bathydoris broadly resembles that of the Dorids, but with much less 
concentration of the ganglia. It lies on the top of the buccal bulb, the cerebral, 
pleural, and pedal pairs being quite separate, but lying close together. As fig. 7 shows, 
the ganglia are asymmetrical in shape and disposition. Each cerebral ganglion (e.g.) 
gives off four nerves from its anterior edge which go to the lips and oral tentacles, 
dividing as they go into a number of smaller nerves. The last bifurcations have small 
ganglionic swellings at the point of division, as in some Tectibranchs. On the posterior 
edge of the dorsal surface of the cerebral ganglion stands a small, almost sessile, 
proximal rhinophorial ganglion (p.rh.g.) which sends a stout nerve (rh.n.) to the dorsal 
tentacle. A distal rhinophorial ganglion swelling (not shown) marks the point of sub- 
division of the rhinophorial nerve as it enters the tentacle. No sub-cerebral commissure 
was found, so that the cerebrals are connected together below the alimentary tube by 
the stomato-gastric loop (s.g.l.) only. The pedal ganglia (ped.g.) are connected by a 
stout pedal (ped.com.) and a slender parapedal commissure (p. ped.com.). In the notch 
between the pleural and pedal ganglia of the right side is placed a small genital ganglion 
(gen.g.) which is broadly united to the pedal and connected with the pleural by a band 
of fibres from its lower aspect. From it four nerves go to the genitalia, both male 
and female, over the surface of which they distribute themselves with a number of local 
ganglionations on their courses. 

The pleural ganglia (pl.g.) give off two main lateral nerves on each side which 
supply the whole of the dorsal integument, with the exception of the anterior region 
which receives a number of very fine nerves from the pleurals not shown in the figure. 
The longer pair of pleural nerves pass back to the anal region, where they anastomose 
with each other and with a visceral nerve (br.n.) from the under surface of the right 
pleural ganglion. From the ganglia on this plexus the gills are supplied as in Doris 
tuberculata. The visceral ganglion, so obvious on the under side of the right pleural 
ganglion of Doris, is not represented as a discrete mass in Bathydoris. The visceral 
ganglion of Doris would appear to include the visceral centres as well as the penial 
centre usually associated with the pedal. In Bathydoris, however, the genital centres 
of the visceral seem to be segregated from the rest and to be associated with the penial 
centre on the pedal to form a special genital ganglion. The long, finely ganglionated 
visceral loop (v.l., v. I.') about the middle of its course sends backwards the chief visceral 
nerve (v.n.), which, after giving a branch to the gastro-oesophageal anastomosis on the 
stomach and liver, continues its course as the reno-cardiac. Two delicate nerves from 
the pleuro-pedal angle (see diagram) supply the branches of the aorta. The stomato- 
gastric ganglia (st.g.g.) are relatively very large, but their size is not surprising when 
we remember the dimensions and muscular complexity of the buccal mechanism which 
they innervate. The inequality in length of the two cerebro-buccal connectives is pro- 
bably the result of the sharp sinistral bend of the oesophagus. There are no separate 



THE ANATOMY OF A NEW SPECIES OF BATHYDORIS. 197 

gastro-oesophageal ganglia, and the paired gastro-oesophageal nerves (g.o.n.) arise 
directly from the stomato-gastric ganglia. The paired nerves pass back in the con- 
nective-tissue investment of the oesophagus to the stomach, where they form a joint 
ganglionated anastomosis with the gastric branch of the visceral, as already described. 
From this plexus the stomach and liver receive their innervation, while the salivary 
glands are supplied from the gastro-cesophageals on their way down the oesophagus. 

This compound system of gastro-hepatic ganglia is paralleled in the Dorids, where 
it was described by Alder and Hancock, while Dreyer has lately shown that an 
analogous arrangement is present in Aeolids and Tritonids. 

Like the other species examined anatomically, B. brotvnii showed no trace of organs 
of sight, and otocysts could not be found by careful surface examination. They were, 
however, found on staining and clearing and also in sections of the brain. They are 
two small sacs placed close to the pleuro-pedal connectives on their lower aspect 
and partially imbedded in the connective -tissue capsule that surrounds the brain. The 
author has found small otocysts similarly placed in some of the Polyceratidae. Otocysts 
were not found in the species examined by Bergh and by Eliot. 

The Excretory System. (Fig. 8.) 

The kidney of Bathydoris is unusually well developed. This was also noted by Eliot, 
who described two fern -like organs lying over the liver as well as the renal syrinx. 
Eliot, however, misconceived the nature of the renal organ, since he took the paired, 
fern-like structures to represent the main portion of the kidney, whereas they are 
merely outgrowths of its floor or ventral wall in the posterior half. The whole renal 
organ is a huge sac extending from almost the extreme posterior end of the body to 
within a short distance of the head, but narrowing in front on account of pressure 
between the alimentary canal and the genital mass. Posteriorly, its delicate dorsal 
wall is overlain by the pericardium, to which it is connected by fibres. Elsewhere it 
bulges free except where it is pinched by the intestinal loop (int.). This dorsal wall 
is throughout non-glandular, except at two points — namely, at its extreme anterior 
corner (h.g.') and at a place in front of the pericardium (b.g.") where a diverticulum of 
it lies as a flap across the intestine. These two points will be further mentioned in 
connection with the vascular system, because the two phagocytic or blood glands lie 
here adherent to the wall of the kidney. The glandular part of the kidney is therefore 
almost entirely confined to its ventral wall and to those parts of it which are folded 
inwards into the renal cavity as the two fern-like structures seen by Eliot. These, 
however, are not two but six in number, the posterior pair being more fern-like than 
the other two which lie on the surface of the gonad (h.g.). These glandular regions 
coincide with the areas of distribution of great branches of the aorta, and the narrow 
strip-like folds forming the two anterior pairs may easily be mistaken for the arterial 
trunks themselves, which actually lie within them below the renal wall. The vascular 



198 MR T. J. EVANS ON 

supply of the kidney is therefore purely arterial, and all the renal arteries arise from 
an aortic bulb (a.e.) opening into the ventricle at the point o.v. in fig. 8, and continu- 
ing forwards as the cephalic artery (cart). The extensions of the renal arteries into 
the gonad are not shown in fig. 8. The blood delivered by the renal arteries — which, 
as we have seen, occupy the crests of the glandular folds — passes on into A^enous lacunae 
which lie deeper in the substance of the folds, and open into a great median venous 
space, lying between the kidney and liver behind, and between the gonad and liver in 
front. Into this median venous space also passes the blood that has traversed the 
gonad and liver. 

The main collecting reservoir of the kidney lies behind the gonad, and in its hinder 
wall is seen the opening into the renal duct which leads to the exterior. In it 
originates the reno-pericardial duct (r.p.d.), consisting of a median tube opening in 
front by a funnel into the renal chamber, and a renal syrinx (r.s.) opening on the floor 
of the pericardium. The syrinx is a bulbous structure with a wide lumen which is 
almost filled with delicate laminate ingrowths of the epithelial lining. Sections of the 
floor of the kidney show that the gland cells lying in connection with the renal arteries 
contain concretions, often of large size, which stain faintly with basic dyes. The con- 
cretions collect in big vacuoles, which finally burst and liberate the excreted contents. 
The foliations of the wall of the syrinx are covered with cells of two kinds. The distal 
part of a lamina — namely, the free edge towards the middle of the lumen — is covered 
by ciliated cells only, the cilia being extremely long. The proximal part — namely, that 
nearer the wall of the syrinx — is glandular, and the cells contain fine granules of a 
substance which takes acid dyes. These cells are continued on to the wall of the 
pericardium. The renal organ of Bathydoris and its vascular supply are thus Dorid 
in type, the reno-pericardial duct, especially, being almost identical in structure with 
its homologue in Doris. The association with the blood glands, non-functional though 
that may be, the absence of ramifications into underlying organs, and its forward 
extension into the head region are features not paralleled among true Dorids. 

The Vascular System. (Figs. 9 and 10.) 

In general, the vascular system of Bathydoris resembles that of the Dorids, but in 
several respects distinct affinities with the blood system of the Pleurobranchids are 
exhibited. It may be conveniently described under the following heads : — 

(1) The Heart. 

The most obvious feature of the heart and pericardium is their asymmetrical 
disposition, since the an tero- posterior axis, unlike that of the true Dorids, lies at an 
angle to the long axis of the body. The pericardium is a spacious cavity lying 
posteriorly on the surface of the kidney, with the reno-pericardial opening in its 
extreme right-hand corner. 



THE ANATOMY OP A NEW SPECIES OF BATHYDORIS. 199 

The asymmetry of position mentioned above is an insignificant matter compared 
with the structural asymmetry shown by the heart itself. The typical Dorid heart is 
roughly an isosceles triangle with three efferent ducts opening into its base, the efferent 
branchial in the middle and the two lateral integumental sinuses at the corners. The 
auricle of Bathydoris, on the contrary, receives but one efferent vessel, which enters it 
at the right-hand side, the efferent branchial vessel and the lateral sinuses being con- 
fluent outside the pericardium altogether, as in the Pleurobranchids. The left side of 
the auricle is fused for some distance with the pericardial wall, along which it sends a 
muscular wing. This asymmetry, as we shall see later, is only one of many pre-Dorid 
and ancestral opisthobranch features exhibited by Bathydoris. 

(2) The Arterial System. 

Although the arterial system possesses no striking feature, it is proposed to describe 
it somewhat fully, because no comprehensive account exists of the arterial system of 
any Dorid except Hancock and Embleton's account of Doris tuberculata in their 
famous article in the Philosophical Transactions of the Royal Society. The ventricle 
(v.) is immediately followed by a large aortic bulb (a.c.) from which arise the renal 
arteries (see kidney), as well as vessels to the intestinal loop, the gonad and the 
periphery of the liver mass lying below. The aortic bulb is continued forwards as the 
main cephalic artery (cart.) This gives off on the left the visceral artery (v.art.), 
running below the intestine and supplying the liver, stomach, and oesophagus. In 
fig. 9 the arterial trunks lying below the outlined viscera are dot-shaded. After 
giving off the genital arteries (g.art.) on the right, the cephalic artery bifurcates, one 
branch passing over the oesophagus to the left and the other below the buccal mass to 
the right. The left branch provides both salivary glands (sal.g.), the brain (cer.art.), 
and the buccal muscles of both sides, while the right branch goes direct into a spacious 
infra-pharyngeal lacuna (lac), in which the left also ends. This lacuna was also found 
in Doris tuberculata. It should be noted that the cephalic artery forms a complete 
collar round the oesophagus and buccal bulb. From the central lacuna under the bulb 
arise a number of vessels. A median vessel passes straight up into the bulb (bucart.) ; 
three run forward into the lips (lab.art.) and floor of the mouth, while a broad median 
vessel dips into the foot and bends backward in its substance, to continue throughout 
its length as a median pedal artery (ped.art.) 

(3) The General Hsemocosle. 

The irregular lacunar blood-space in which lie all the viscera is in Bathydoris 

nowhere spacious. It receives the blood that has passed through the tissues from the 

arteries, except the renal, gonadial, and hepatic blood, which is collected in another 

way already indicated in the description of the kidney. The hsemocoelic blood passes 

partly into the gills and partly into the dorsal integument, but the proportion of blood 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 6). 27 



200 MR T. J. EVANS ON 

that passes into the integument is much in excess of that in the Dorids. The thin 
skin, studded with papillae, is conducive to this amplification of the tegumentary 
respiration in Bathydoris, while the thick, glandular, and spiculose skin of the Dorids 
has vitiated this system and necessitated the extension of the special gills. The dorsal 
wall of the haemoccele is a membrane more or less bound to the underlying organs by 
conjunctive-tissue fibres. This membrane is separated from the dorsal integument by 
an empty space, but runs into it at the side of the body all round. Thus, when an 
incision is made through the dorsal body- wall, the space entered is not the haemoccele, 
but this closed cavity between the body-wall and the dorsal wall of the haemoccele. 
The same arrangement is present in the Dorids alone among Nudibranchs, though the shell- 
cavity of Pleurobranchus closely resembles the problematic dorsal cavity of the Dorids. 
Whatever be the nature of the cavity, Hancock and Bmbleton's name — peritoneum 
— for its lining should not be perpetuated. The passage of blood from the underlying 
haemoccele into the skin and its papillary outgrowths takes place below the level of the 
edge of the dorsal hsemocoelic wall all round. The blood that runs from the haemoccele 
to the gills passes along two narrow conduits on the posterior aspect of the liver (h.v., 
fig. 10). This must be regarded as of secondary importance in the afferent branchial 
system. 

(4) The Afferent Branchial System. (Fig. 10.) 

Blood enters the branchiae from two sources : (a) from the haemoccele by the small 
paired ducts (h.v.) already mentioned, and (b) from a great median venous space (m.s.) 
lying above the liver, which receives the blood from the kidney, liver, and gonad. Just 
before narrowing in order to enter the gills (a.b.v.), it receives the paired ducts from 
the haemoccele (a). (See also the description of the kidney.) The afferent space at the 
base of the gills is not a circle, as in the Dorids, but a transverse expansion of the 
afferent vein from which ramifying tubes run up the branchiae. 

(5) The Efferent Branchials. 

The afferent and efferent venules in the gill-leaflets form continuous loops from the 
afferent to the efferent side of a gill lobe, and the efferent veins from the two gills join 
together to form a transverse space at the base of the gills similar to the contiguous 
afferent space. This space is connected with the auricle by a tube (e.b.v.) running to 
the right and entering the auricle at its right-hand corner. 

(6) TJie Efferent Tegumentary System. 

The blood that enters the skin and its papillae from the haemocoele returns from all 
sides into a circular sinus (c.s.) running round the edge of the pericardium. The 
efferent tubules returning blood into the sinus were described by Bergh as renal tubules 
in B. abyssorwm. The sinus opens behind into the efferent branchial vein just before 



THE ANATOMY OF A NEW SPECIES OF BATHYDORIS. 201 

it reaches the right corner of the pericardium to empty itself into the auricle. The 
circular sinus is also represented in the Pleurobranchids, where it also opens into the 
efferent vessel, but takes a wider sweep round the body. In the Dorids it is repre- 
sented by the two lateral sinuses, which there, however, enter the auricle separately 
and directly. 

(7) The Blood Glands, (b.g.' and b.g." Figs. 8 and 9.) 

The structures commonly called " blood glands " are characteristic of some Tecti- 
branch families (Bullids and Pleurobranchids) and of the Dorids. They are lymphatic or 
phagocytic glands situated on the course of the cephalic artery and supplied by it. In 
the Dorids they lie in the head region near the brain, but in the Tectibranchs they are 
placed further back. In Bathydoris they form two separate masses united to the wall 
of the kidney. It is of some interest to note that in the Prosobranchs possessing them 
they are also associated with the kidney, so that in this, as in many other respects, 
Bathydoris presents features more primitive than the true Dorids, the equivalents of 
which are found among Tectibranchs rather than Nudibranchs. As already mentioned, 
Bergh and Eliot confused the unusual salivary glands with the blood glands, the 
identity of which they did not recognise. 

The Respiratory System. 

As already indicated in connection with the vascular system, the general pallial 
respiration is rendered more effective by the papillary outgrowths, and the blood so 
oxygenated returns into the circular sinus. The special respiratory organs or branchiae 
are two tufts placed symmetrically on a transverse ridge in front of the anus. Each 
tuft stands on a broad base or stalk in such a way as to give the appearance of a roughly 
pinnate condition. The lobate units of the tuft resemble the pinnae of the gill of 
Pleurobranchsea, being laminate on opposite faces of a wide rhachis, while the afferent 
and efferent vessels occupy its edges. The laminae vary in size from mere ridges across 
the face of the rhachis to longish leaves which may themselves be provided with laminae. 
In this way an irregular bipinnate condition is simulated. It will be seen that, by 
narrowing the rhachis so as to bring the ascending and descending vessels nearer 
together and regularising the pinnation, the Dorid plume would be produced. On the 
other hand, if the laminae were equal in size and the tufts stretched along the ridge in 
a regular row, the sessile portion of a Pleurobranchid ctenidium would result. The gill 
of Bathydoris brownii would therefore appear to be in a condition intermediate between 
a typical Dorid rosette of plumes and a Tectibranch gill. There is, however, no indica- 
tion of a circumanal circlet either in the gill itself or in the underlying vessels, and though 
the tufts are provided with muscles capable of reducing their height, they cannot be 
retracted below the general level of the integument. 



202 MR T. J. EVANS ON 

The Reproductive System. (Fig. 12.) 

No adequate description or figure of this system in a Bathydoris has hitherto been 
given, but both Eliot and Bergh give a somewhat hesitating impression that it is 
constructed on the Dorid plan. Since the universal triauly of known Dorids is one of 
their most striking characteristics, it is essential that on such a critical point our 
knowledge should be clear and definite. 

The hermaphrodite gonad (h.g.) lies posteriorly below the kidney and above the liver 
within the arc made by the intestine. It is a yellowish, bi-convex lenticular body, 
truncated in front and with a minutely lobulated surface. The specimen was captured 
at the stage of male activity in the protandric cycle, since the male acini and ducts 
are full of sperms, while the eggs are small and lightly yolked. Its blood supply is 
an extension of the renal arterial system, branches from which pass through the lower 
wall of the kidney into its substance. 

The common hermaphrodite duct (c.h.d.) leaves the gonad as a single slender 
tube. It is ampullated (amp.), as usual in Nudibranchs, but its extreme length escaped 
previous notice. After a short, slender portion beyond the ampulla, it divides into 
two tubes, the vas deferens (v.d.) and the oviduct (o.d.). The vas deferens is a 
comparatively short, coiled tube, expanded by the glands in its walls into a prostate 
for nearly the whole of its length. It enters the penis sac (p.s.) some distance 
from the end and runs a straight course to the tip of the everted penis imbedded 
in loose connective tissue and muscle fibres. The mode of extroversion of the 
penis, deducible from dissection of the everted organ, is represented in the section- 
diagrams (fig. 11, a and b), the dotted area representing loose fibrous tissue the 
perfusion of which with blood from the hsemocoele causes the extroversion. The penis 
is seen to be a partial introvert, since the end is retracted into the sac unchanged. 
This terminal portion (p.) presents a remarkable appearance on account of the sucker- 
like pits covering one side of it. It is possible that the pits, under control of the 
blood-pressure in the penis, really act as suckers on the smooth surface of the female 
atrial wall. 

The oviduct soon enters the massive mucus-albumen gland complex (m.g. and a.g.), 
the structure of which could not be investigated on account of its stony hardness. The 
albumen gland could be recognised on the upper surface by its yellowish-brown colour 
and its granular consistency. The coils of the mucus gland end distally as the broad 
tube which opens into the female atrium. The atrium is turned out as in copulation : 
the first part of it has a highly crinkled surface, but inside this arise two leaf-like lips 
(a.l.), or folds of its surface, which between them enclose the entrance (f.o.) into the 
female channel. These valvular lips appear to be a characteristic feature of Bathydoris, 
because they are also partially shown in surface view in B. clavigera by Thiele. 
Within the valve on the posterior wall of the channel opens the vagina (vg.), which 
consists of a stout tube ending blindly in a globular recurved bursa copulatrix (b.c). 



THE ANATOMY OF A NEW SPECIES OF BATHYDOFJS. 203 

There is no second sac on the course of the vagina, which is also the only connection 
between the bursa and the female channel. From this it follows that the herma- 
phrodite duct of Bathydoris divides but once — namely, into a male and a female duct. 
The Dorid duct bearing the so-called spermatocyst, and connecting the bursa with the 
region of fertilisation in the course of the oviduct, is totally unrepresented. In short, 
Bathydoris brownii is typically diaulic, like Tritonia and the Pleurobranchids. 

Externally, the everted organs are surrounded by a rim representing the limit of 
the common genital vestibule of the male and female system. 

Synopsis of Specific and Generic Characters. 

When we come to analyse the differences between the foregoing account and those 
of the two previous investigators of the genus, we encounter considerable difficulties in 
deciding which are specific differences and which may be put forward as corrections. 
To the latter category we may presume to relegate all differences in regard to which 
previous statements have been qualified or made with reservation. 

The specific distinctness of Bathydoris brownii is undoubtedly more striking than 
that of any of the other recorded species, and, since the animal was mature, there can 
be no question of its being the young of any one of them. Among characters pre- 
sumably not of generic rank which distinguish the species may be mentioned the 
following : — 

( 1 ) The uniformity in size and shape of the lateral teeth of the radula. 

(2) The pitted pad on the glans penis. 

(3) The immediate sinistral bend of the oesophagus at its origin from the buccal 

mass and the unequal lengths of the cerebro-buccal connectives. The 
causal connection between these two features stamps them as true anatomical 
constants. 

(4) The two tufted gills placed on a traverse ridge symmetrically in front of the 

anus. 
On the basis adopted above, it is likely that the following features in which the present 
account differs from those of Bergh and Eliot may be regarded as of generic value : — 

(1) The diauly of the reproductive system. 

(2) The circular canal embracing the pericardium and collecting blood from the 

dorsal integument. 

(3) The follicular nature and great size of the salivary glands. 

(4) The presence of two blood glands on the wall of the kidney (the structures 

described as such by Bergh and Eliot turned out, on histological examina- 
tion, to be the follicles of the salivary glands ; the true blood glands were 
not seen by these authors). 

(5) The asymmetrical opening of the efferent vein into the auricle. 

(6) The great saccular kidney with its ventral wall thrown into folds. 



204 MR T. J. EVANS ON 

(7) The segregation of the genital elements of the visceral into a distinct ganglion 

on the surface of the brain. 

(8) The possession of a proximal and a distal rhinophorial ganglion. 

(9) The cuticularisation of the wall of the alimentary canal as far as the stomach 

(the horny cones imbedded in the cuticle were described by Eliot for B. 
hodgsoni, but no armature was found by Bergh in B. abyssorum and B. 
ingoljiana ; its systematic value cannot therefore be assigned). 

(10) The presence of a gastric csecum. 

(11) The presence of small otocysts below the pleuro-pedal connective. 

The genus may now be defined in the following terms, of which some are supple- 
mentary to Bergh's original definition : — 

Body highly arched and elliptical in outline. The edge of the notseum slight or 
wanting. Dorsal papillae present or absent. Ehinophores placed rather far back, non- 
retractile, perfoliated. Gills in front of the anal papilla, variable in number, non- 
retractile. Buccal mass very bulky. Radula sac not an appendage. Dental formula 
n. 1. n. Buccal cavity with a thick cuticle extending down the oesophagus. Powerful 
jaws present. (Esophagus may have horny cones. Liver massive and unlobed, not 
invaded by any other viscus. Salivary glands follicular, flattened, with a stout 
duct. Cerebral and pleural ganglia distinct. Stomatogastric loop very long. No 
gastro-cesophageal ganglia, but the long gastro-oesophageal loop arises from the 
buccals. Eyes absent. Kidney saccular with laminate ingrowths of its ventral wall. 
Branchial and pallial efferents join before entering the right side of the auricle. Penis 
unarmed and massive. Hermaphrodite gland a compact mass. Reproductive system 
diaulic. 

The Affinities of Bathydoris. 

Bergh and Eliot have invested Bathydoris with a certain importance as a type 
combining the features of the Dorids with certain Tritonid characters, with prepon- 
derating affinities to the Dorids. The Tritonid features accentuated by Bergh were 
the buccal apparatus and the unarmed penis, while Eliot rightly passes over the latter 
resemblance unnoticed, since an armature of the penis may be present or absent among 
the species of some genera of Dorids. As to the buccal apparatus, even a superficial 
examination shows that, when reference has been made to the great size of the buccal 
muscles and the jaws, the sole resemblance has been stated in full. In Tritonia the 
odontophoral mass arises from the dorsal wall of the buccal cavity and bulges down- 
wards, while in Bathydoris that organ arises from the floor of the buccal cavity and 
bulges upwards. The mandibles are also quite differently placed and used in the two 
animals, those of Tritonia having their long, finely serrulated cutting edges facing the 
floor of the mouth cavity below the radula mass, while the blunt beaks of the mandibles 
of Bathydoris jut into the mouth above the radula. Moreover, the oesophagus takes 
its origin on the hinder aspect of the globular buccal bulb of Bathydoris, while the 



THE ANATOMY OF A NEW SPECIES OF BATHYDORIS. 205 

oesophagus of Tritonia rises out of the dorsal surface of the bulb and well forward, the 
main bulk of muscles being behind it. These and corresponding differences in the 
muscular mechanism suffice to make good the statement made above, that the supposed 
resemblances are confined to size and the presence of powerful jaws. It should be 
mentioned that Bergh refers specially to Bornella in assigning Tritonid features to 
Bathydoris ; without discussing the problematic relationship of Bornella to the 
Tritonids, suffice it to state that the large buccal apparatus of that form differs from 
that of both Tritonia and Bathydoris. 

It is indeed likely that these three cases of powerful and mandibulate mouth parts 
are examples of convergence in unrelated types. The only other reference to a non- 
Dorid affinity of Bathydoris is made by Eliot when he compares the armature of the 
"stomach" with that found in Bornella. This comparison is strange, coming from an 
author who has since, in the Ray Society's monograph, separated the two genera in 
his first cleavage of the Nudibranchs. In any case the comparison is untenable, since 
the two armatures are totally unlike in structure and position, that of Bornella being 
situated in a region of the alimentary tube posterior to the point of entrance of the 
liver ducts. On the foregoing grounds we must regard the proposed Tritonid and 
Bornellid affinities of Bathydoris as inadmissible. It is, however, obvious that the 
investigation of this last species has brought out certain features of the genus which 
render necessary the reopening of the discussion of its affinities and, as we shall see 
later, those of the Dorids generally. It is no less certain that the genus presents a 
combination of characters far more significant than that considered by Bergh when he 
assigned its affinities — namely, a Dorid gill of a primitive form, an asymmetrical heart 
and efferent branchial system, blood glands placed far back on the course of the aorta, 
a thin integument with scattered branchiate outgrowths, a diaulic reproductive system, 
a liver distinct from the gonad and kidney, a brain with separate ganglia, a nerve 
collar embracing the buccal bulb and not the oesophagus, and finally, but perhaps least 
significant, a powerful buccal apparatus. 

That Bathydoris must be definitely placed among doridiform animals follows from 
its possession of the following striking Dorid characteristics : — 

(a) The collocation of the anus, renal pore, and gills in the median line posteriorly. 

The gill is, however, more primitive than the typical rosette form common 
among Dorids, though primitive gills are also found in such types as 
Trevelyana and Nembrotha. 

(b) With the exception of the buccal mass and the protected oesophagus, the 

alimentary canal is Dorid, even to the possession of a gastric ceecum, and 
those divergent features are adaptations to a coarser and more omnivorous 
diet. The enlargement of the salivary glands is probably due to the same 
cause. 

(c) The kidney is a Dorid structure, the reno-pericardial tube and syrinx being 

practically identical with those of Doris as described by Hancock 



206 MR T. J. EVANS ON 

and by Hecht. The absence of ramifications is, doubtless, a primitive 
character. 

(d) The blood system is built essentially on Dorid lines, but presents a greater 

number of primitive features than any other. Chief among them are the 
possession of but one auricular efferent opening, the union of the circular 
collecting canal of the integumental system with the efferent branchial, and 
the position of the blood glands. It is noteworthy that these primitive 
features are points of agreement with the Tectibranchs, especially the 
Pleurobranchids. 

(e) As to the reproductive system, its diaulic condition makes it more primitive 

than that of any other known Dorid ; but, apart from that very important 
divergence, it closely resembles that of Doris, since the separation of the 
gonad from the liver is found in a typical Dorid like Alloiodoris. 

(f) The nervous system, in spite of a close similarity to the Dorid type in most 

respects, differs from it in several important points. Of these, the length 

of the nerve collar and the position of the brain on the top of the buccal 

mass are paralleled in Tritonia and the Pleurobranchids, and should probably 

be regarded as primitive, while the distinctness of the ganglia of the brain 

and the absence of separate gastro- oesophageal, if primitive features, take 

us back to a condition earlier than that found in the Pleurobranchids and 

Tritonia. The fusion of the ganglia of the visceral loop with the pleurals 

is, on the other hand, a modern feature, as is the loss of eyes consequent 

on the adoption of a deep-water habitat. 

We conclude, therefore, that Bathydoris is a highly primitive Dorid possessing 

some characters that adapt it to a specialised habitat and mode of life, while those that 

are primitive connect it with the Tectibranchs, particularly the Pleurobranchids among 

existing forms. The derivation of the Dorids from Pleurobranchid ancestors is, 

however, no new proposition. Guiart, for example, has recently advocated their union 

into one group, and Pelseneer has derived all Nudibranchs from the Pleurobranchids 

with Tritonia as an intermediate link. 

Bergh's advocacy of a special relationship between Bathydoris and Tritonia on the 
evidence of the buccal apparatus has already been criticised. Pelseneer's position, 
however, takes a wider outlook, but takes no cognisance of Bathydoris at all. He 
bases his contention of the Tritonid origin of all Nudibranchs on the possession by 
Tritonia, in common with the Pleurobranchids, of a large number of primitive Nudibranch 
characters which are not found together in any other Nudibranchs. These are : — a 
frontal veil, formed by the fusion of the oral tentacles of the Pleurobranchid, a wide 
foot, a ventricle turned to the right, a broad raclula, a nervous system placed on the 
buccal bulb, an oesophageal crop, extensive salivary glands, a saccular, unramified 
kidney, a long reno-pericardial tube, pericardial glands on the auricle, male and female 
openings in a common vestibule, and a lateral anus. Of these, it will be noticed that 



THE ANATOMY OF A NEW SPECIES OF BATHYDORIS. 207 

Bathydoris possesses all except the lateral anus. Further, it retains oral tentacles in 
a condition more strongly reminiscent of those of Pleurohranchsea than that of the oral 
veil of Tritonia, while the separate ganglia of the brain and the separate gonad of 
Bathydoris can certainly not be regarded as new and derived features in that genus. 
Thus, excluding the case of the lateral anus, which will be considered separately, Bathy- 
doris, which is essentially Dorid in construction, exhibits all the primitive features of 
Tritonia, some indeed being more primitive than the corresponding ones in Tritonia, 
the supposed ancestor. At this reductio ad absurdum we arrive by considering only 
those primitive features selected by Pelseneer, without calling in the evidence of the 
blood and respiratory systems wherein Tritonia, with its symmetrical auricle receiving 
blood from symmetrical lateral sinuses, appears very modern indeed. It is in the complete 
avoidance of any comparison between vascular and respiratory systems in Tritonia and 
Pleurobranchids that the weakness of the Tritonid theory of Nudibranch descent lies, 
and it is significant that on the characters of these very systems is primarily based any 
discussion of gastropod and even molluscan affinities. Previous application of this 
criterion in the Opisthobranchs has resulted in their cleavage into Tectibranchs with 
a ctenidial gill and Nudibranchs with pallial outgrowths of varied form and distribution 
replacing the lost ctenidium. Of these neomorphic gills the lateral tufts of Tritonia 
have been regarded as an early type, but it is not clear whether the Dorid circlet was 
derived from them by concentration or by local specialisation round a posterior anus or 
was evolved independently. Nor is it clear why modern writers on the Opisthobranchs 
have always accepted the neomorphic nature of the Dorid circlet. It is true that a 
comparison of the highly specialised, multipinnate plumes placed in a pit in the 
tuberculate dorsum of some Dorids provides no suggestion of homology with the 
ctenidium of a Tectibranch ; but it is not such a comparison of extremes that evinces 
homologies. In Bathydoris, however, the gill is in two portions only, joined by a 
crinkled ridge, it shows but the beginnings of pinnation, its lobes have the broad laminse 
running from the afferent to the efferent side seen in the ctenidium of the Tectibranch, 
and there is no suggestion of the circumanal ring in either the gill or the underlying 
vessels. From this point of view the extreme similarity of the condition of the 
auricle, the efferent branchial vessel, and the circular sinus in Bathydoris and the 
Pleurobranchids acquires a special significance. Evidence derived from the nature of 
the innervation is perhaps of doubtful value ; but, so far as it goes, it is favourable 
to the present contention, since the Dorid gill is jointly innervated from pleural and 
visceral centres, while other Nudibranch gills receive no visceral nerves unless invaded 
by ramifications of the liver. The dorsal position of the Dorid gill should present little 
difficulty, since the pallial edge of the Dorids is undoubtedly a new formation of 
mechanical value which progressively increases in width within the group and is absent 
in many genera. In any case, the same difficulty would apply to the anus and renal 
pore, and there is no proposal to class them as new formations in the Dorids. The separa- 
tion into two or more parts also forms no objection to the ctenidial nature of the Dorid 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 6). 28 



208 MR T. J. EVANS ON 

gill, since it is a progressive process in the group and is incipient in many Tectibranchs, 
including Aplysia. Moreover, at least one Dorid, Trevelyana crocea, has a single un- 
divided laminate gill indistinguishable from a ctenidium. Furthermore, the three residual 
units of the pallial complex — namely, the kidney and its pore, the intestine with the anus, 
and the gill — occupy in the Dorids just those relative positions which they would occupy 
had they been turned over from the Tectibranch position into the median dorsal line. 
Here one is inclined to ask what is the nature of the great cavity, cut off from the 
underlying hgemoccele, which lies under the dorsal integument of the Dorid, but is 
absent in all other Nudibranchs. An exactly similar cavity in Pleurobranchus or 
Oscanius contains a shell-remnant, and is the shell-cavity. In the absence of any 
information regarding the metamorphosis of the veliger of either Dorids or Pleuro- 
branchids, it is difficult to find any satisfactory reason for contradicting the homology 
of these two spaces. 

It is on the above grounds proposed to define the Dorids as ctenidiate Opisthobranchs 
that have retained the shell-cavity and in which the elements of the pallial complex 
have moved dorsally into the median line. In this position the ctenidium has under- 
gone progressive modification within the group, the retractile circlet being its highest 
development. 

In Tritonia, on the other hand, the residual members of the pallial complex have 
remained in a more anterior position than they occupy in many Tectibranchs, and in 
that position the old molluscan gill has been lost. Whereas in the Dorids and Pleuro- 
branchids the connection of the auricle with lateral integumentary sinuses is supple- 
mentary to the ctenidial connection, in Tritonia it is the sole remaining connection of 
the auricle with respiratory sinuses. As a primitive actenidiate animal, however, 
Tritonia retains many common features with the Dorids and Pleurobranchids. its 
nearest ctenidiate relatives. 



BIBLIOGRAPHY. 



(1) Bergh, R., "Report on the Nudibranchiata," " Challenger" Expedition, vol. x., 1884. 

(2) "Nudibranchiate Gasteropoda," Danish "Ingolf" Expedition, vols, ii.-iii., 1900. 

(3) Dreyer, T. H., "tjber das Blutgefass- und Nervensystem der Aeolididae und Tritoniadae," Zeitschr. 

f.wiss. Zool., Bd. xcvi., 1910. 

(4) Eliot, Sir Charles, A Monograph of the British Nudibranchiate Mollusca, part viii., Ray 

Society, 1910. 

(5) " Report on the Mollusca Nudibranchiata collected by the Discovery," National Antarctic 

Expedition Reports, 1907. 

(6) Guiart, J., " Les Mollusques Tectibranches," Causeries scientifiques de la Soc. Zool. de France, 1900. 

(7) Contribution a I'etude des Gasteropodes Opisthobranches, Lille, 1901. 

(8) Hancock, " On the Structure and Homologies of the Renal Organ in the Nudibranchiate Mollusca," 

Trans. Linn. Soc. Lond., xxv., 1865. 

(9) Hancock and Embleton, "On the Anatomy of Doris," Phil. Trans., London, 1852. 

(10) Hecht, E., "Contribution a 1'dtude des Nudibranches," Mem. Soc. Zool. de France, vol. viii., 1895. 



THE ANATOMY OF A NEW SPECIES OF BATHYDORIS. 209 

(11) de Lacazk-Duthiers, H., "Histoire anatomique et physiologique du Pleurobranche orange," Ann. 

Sci. Nat. (4), xi., 1859. 

(12) Moquin-Tandon, G., Recherches anatomiques sur VOmbrelle de la Mediterranee, These de Paris, 1870. 

(13) Pelseneer, P., Recherches sur divers Opistobranches, Gand, 1893. 

(14) Thiele, J., "Die antarktischen Schnecken und Muscheln," Deutsche Sudpolar Expedition, 

Berlin, 1912. 



EXPLANATION OF PLATES XVII. AND XVIII. 

Bathydoris broionii. 

Fig. 1. The animal seen from the dorsal side, natural size. 

Fig. 2. General view of the alimentary system from above, b.m., buccal mass ; cr., oesophageal crop ; 
int., intestine; I., liver; oe., oesophagus; p.m., lateral pads of the inner lips; s.g., salivary glands. 

Fig. 3. Buccal cavity laid open from above. J., jaws ; o., odontophore ; 03., oesophagus ; P. J., pads of 
the jaw ; p.m., pads of inner lips ; R., radula. 

Fig. 4. Portion of a half-row of the radula. L. 1 -L. T , lateral teeth 1 to 5 ; Rh., rhachidian tooth. 

Fig. 5. Stomach and adjacent parts of the alimentary canal laid open ; the cut is continued into the 
posterior lobe of the liver, b.o., bands on the wall of the crop ; I., liver ; int., intestine ; l.t., liver ducts; s.r., 
gastric caecum ; st., stomach. 

Fig. 6. Section across an oesophageal band, b.c, brown cones ; ct., cuticle ; ep., epithelial layer ; m., 
muscle layers. 

Fig. 7. Nervous system, b.co., buccal commissure ; e.g., cerebral ganglion ; gen.g., genital ganglion ; 
g.o.n., gastro-cesophageal nerves ; ped.g., pedal ganglion ; ped.com., pedal commissure ; p.ped.com., parapedal 
commissure ; pl.g., pleural ganglion ; s.g.l., buccal loop ; st.g.g., stomato-gastric or buccal ganglion ; v.l., v.l. 1 , 
visceral loop ; v.n., visceral nerve. 

Fig. 8. Kidney with thin dorsal wall removed, a.c, aortic bulb ; b.g}, b.g. i[ , lobes of the kidney to which 
the blood glands are attached; cart., cephalic artery; h.g., hermaphrodite gonad; int., intestine; o.v., 
opening of aortic swelling iuto the ventricle; r.d., renal tube to exterior ; r.p.d., reno-pericardial duct; r.s., 
renal syrinx. 

Fig. 9. Arterial system, a.c, aortic bulb; b.g}, h.g. [i , blood glands; cart., cephalic artery; cer.art., 
cerebral artery; bucart., buccal artery; g.art, genital artery; lab.art., labial arteries; lac, lacuna under 
buccal mass; ped.art., pedal artery; sal.g., salivary gland; v.art., visceral artery. 

Fig. 10. Diagram of the afferent and efferent vessels, a.b.v., afferent branchial vein; aur., auricle; 
cs., circular sinus ; e.b.v., efferent branchial vein; h.v., haemoccelic vessels; m.s., median sinus. 

Fig. 11, a and b. Diagram showing the relation of the penis to its sheath in the retracted and protruded 
condition. 

Fig. 12. General view of the reproductive system. a.L, atrial lips ; a.g., albumen gland; amp., ampulla ; 
b.c, bursa copulatrix ; c.h.d., common hermaphrodite duct ; f.o., female opening ; h.g., gonad ; m.g., mucus 
gland; o.d., oviduct; p., pitted pad on penis; p.s., penis sac; v.d., vas deferens; vg., vagina. 



is. Roy. Soc. Edin r . 



Vol. L Plate X^ll 



Evans: New Species of Bathydoris. 





g.o.n. 




I RITCHIE*.. 



Vol.L Plate XVIII. 



Evans: New Species of Bathydoris 




■ 



( 211 ) 



VII. — Rupture Stresses in Beams and Crane Hooks. By Angus R. Fulton, 
B.Sc, A.M.Inst.C.B., Engineering Department, University College, Dundee. 
Communicated by Professor A. H. Gibson, D.Sc, A.M.Inst.C.E. 

(MS. received November 4, 1913. Read February 16, 1914. Issued separately April 15, 1914.) 

I. Bending Moment as in a Beam. 

In connection with a research into the failure of timber under stress, carried out 
by the author, and on which a paper was read before the Royal Society of Edinburgh 
and published in their Transactions* many of the tests were made by cross-breaking. 

The probable development of the stress diagram of a beam of rectangular section, 
supported at two ends and loaded in the middle where the tensile and compressive 
strengths of the material are of different values, was illustrated in fig. 38 of that 
paper. It was there laid down that when the fracture point was ultimately reached 
either (a) the tension and compression stress areas both assume the rectangular 
form and are equal to each other, the stress ordinates respectively being equal to 
the ultimate breaking stresses of the material in direct tension and compression ; or 
(b) the cohesion between adjacent fibres measured from the neutral axis outwards is 
not sufficient to withstand the shear induced by the resisting moment of the beam, 
a shear which is at a maximum along the neutral axis of the beam. 

These assumptions made for the case of beams of rectangular sections might be 
said to have found verification in the experiments on such beams of various timbers 
supported at two ends and loaded in the middle, the results of which were tabulated 
in that paper. 

Experiments on Cast-iron Beams. 

Since then the author has carried out a number of similar tests on rectangular 
beams made of cast iron, a material which differs from timber in that it apparently 
possesses no plastic stage such as characterises the latter. 

The specimens used for the purpose of obtaining direct tensile and compressive 
stresses were cast from the same melting as the beams and so were directly comparable. 

The average values obtained for these direct stresses were : — 

Cast Iron. 
Tension stress, T= 9*5 tons per sq. in. 

Compression stress, C = 45 '5 ,, ,, ,, 

The dimensions of test pieces used in compression were 2-f in. long and 1^ in. 

xf in. 

The formula as derived in the paper referred to, and presumably applicable to all 

* Trans. R.S.E., vol. xlviii., 1912, pp. 417-440. 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 7). 29 



212 



MR ANGUS E. FULTON ON 



rectangular beams in which an ultimate rectangular distribution of stress is reached, 

• (1) 



gives : 



b<P C ' bd? T 



where t = the ultimate extreme tensile fibre stress in beam, 

c= „ ,, ,, compressive fibre stress in beam, 

M= ,, external bending moment, 
T= ,, ultimate direct tensile stress of the material, 
C = ,, ,, ,, compressive stress of the material, 

b = ,, breadth, and d = depth of the beam. 

Substituting for M = — — , since the beams were loaded in the centre and supported 

at the two ends, and for T and C the average values obtained, we have 

WL WL 

£ = •70 — - and c=3'33 — - for cast iron. 
bd 2 bd? 

Table I. gives the results of the beam experiments, and it will be seen that the 
calculated values of the extreme fibre stresses t and c agree very well with the 



Table I. 

Cast-Iron Beams. 













Tons per sq. in. 


Elastic Formula. 


xlO 7 . 


No. 


Span. 


b. 


d. 


Breaking 
Weight 




























in Tons. 


*=-70^. 


c = 6 66 — - . 


t = c= 1-5 — w - 


E s . 


E„. 












bd 2 


bd 2 


bd 2 






1 


36 


2-00 


1-02 


•8 


9-7 


46-17 


20-77 


1-48 




2 


12 


1-02 


200 


4-43 


9-14 


43-50 


19 


57 






3 


36 


100 


1-95 


1-45 


9-65 


45-93 


20 


67 




1-85 


4 


12 


1-00 


1-95 


4-29 


9-45 


45-00 


20 


25 






5 


12 


1-95 


1-00 


217 


9-31 


44-33 


19 


95 






6 


36 


1-05 


2 05 


1-6 


9-17 


43-67 


19 


65 




1-53 


7 


12 


105 


205 


434 


8-30 


39-50 


17 


77 






8 


12 


2-05 


1-05 


2-2 


8-61 


41-10 


18 


45 






9 


36 


203 


1-07 


•65 


7-07 


33-67 


15 


15 


1-35 




10 


12 


2-03 


1-07 


193 


7-00 


3333 


15 


00 






11 


12 


107 


2-03 


4-43 


8-47 


40-33 


18 


15 






12 


36 


2-00 


1-05 


•72 


8-26 


39-33 


17 


7 


1-48 




13 


12 


2-00 


1-05 


21 


8-01 


3817 


17 


17 






14 


12 


1-05 


2-00 


4-4 


8-75 


4167 


18 


75 






15 


36 


1-03 


2-05 


1-34 


7-77 


3700 


16 


65 


1-51 


1-51 


16 


12 


103 


2-05 


4-74 


9-18 


43-87 


19 


74 






17 


12 


2-05 


1 03 


243 


941 


44 83 


20 


17 






18 


36 


105 


204 


1-55 


8-94 


42-57 


19 


15 


1-48 


1-67 


19 


12 


2-04 


105 


233 


8-71 


41-47 


18 


66 






20 


12 


1-05 


2-04 


4-75 


9-14 


43-50 


19 


57 






21 


36 


2 06 


1-03 


•76 


8-50 


4050 


18 


22 


1-45 


1-61 


22 


12 


2-06 


103 


2-38 


8-88 


42-30 


19 


03 




• • • 


23 


12 


1-03 


2-06 


4-32 


830 


39-53 


17-79 







RUPTURE STRESSES IN BEAMS AND CRANE HOOKS. 



213 



ultimate stresses obtained in direct tension and compression. The last two columns 
of the table give Young's modulus when the beam is placed on its broad or narrow 
face. 



^ 



~y 



+ 



\ 



General Case for Beams of any Section, but with the Ultimate Distribution 

of Stress a Rectangular One. 

The general case may be deduced as follows : — 
Let fig. 1 denote the section of the beam, and let 
A = total sectional area, 

A t = ,, area ultimately subjected to tensile stress, 
A c = ,, ,, ,, ,, compressive stress. 

t, c, T, C, and M have the same meaning as before. 

Also let NN = ultimate position of neutral axis, the plane 
of which extends along the length of the 
beam and divides each cross-section of the 
beam into areas A { and A e , 
y t = distance of centroid of area A t from NN, 

yc 11 ii ii -"-e 5 > 

and let the ultimate distribution of stress be a rectangular one. 

From the condition (a) that the tension and compression stress areas are equal 

to each other, we have 

tA t -cA e =0. ..... (la) 

If the ultimate fibre stresses reached at the point of rupture be equal to the 
ultimate direct stresses T and C in tension and compression respectively, then 

TA«-CA c = 0, 

and 

A,_C 
A C ~T 

and similarly 




E'&l 



A, + A c T + C 



.-. A, = A 



C 

T + C 



A, = A 



T + C 



We therefore obtain the position of the neutral axis of the beam at fracture point 

by dividing the total area A into areas A { and A„ in the ratio of =. 

Further, since the resisting moment of the section just up to the point of rupture 
equals the external bending moment, 



C T 

-. M = tk t y t + rA c y c = tA t (y t + y c ) = cA c (y t + y c ) = tA— — (y, + y c ) = cA^—^{y t + y c ) 



T + C v 



T + C x 



t=- 



and c = 



^1 T + C 

Hy t +y c ) C 

M T + C 

Hl/t + Vc) T T~ 



(I) 

(La) 
(Lb) 



214 



MR ANGUS R. FULTON ON 



If T and C are equal, the final position of the neutral axis will divide each section 
of the beam into two equal areas and the value of the fibre stress 

t or c = . . . . . ( 1 ) 

In such a case, if the section is symmetrical about the original neutral axis, the 
position of the latter does not change. 

Experiments. 
In order to verify the correctness of such a formula, some tests were made by 
the author on wrought-iron bars. These bars were stressed as beams, and since the 





D 



G 



H 



Eigrl£ 



material used here was ductile, no definite rupture took place. The ultimate load 
was therefore taken as that load at which the beam failed to stand up to the 
applied load and buckled up. Portions were cut out and tested to failure in 
direct tension. No tests, however, were made in direct compression, the fracture 
value of the material under compression being assumed to be equal to that under 
tension. 

The bars were of various sections, and are illustrated in fig. la. 
The sections A and B, which are circular, one solid and the other hollow, show 
an entire agreement between the direct tensile stress and the induced tensile stress. 



RUPTURE STRESSES IN BEAMS AND CRANE HOOKS. 



215 



Of the rectangular specimens C and D agree fairly well, but E, which is square, falls 
short in the induced stress. 

F, G, and H are comparable as a group, and the results in the cases F and Gr are 
quite satisfactory. H, however, shows a deficiency in the induced stress which is 
somewhat marked on the 36-inch span, but less so on the 15-inch span. This may 
be explained by the fact that it is impossible for the web portion of the section as it 
is laid to be stressed to its full rupture value before the flanges give way. But if we 
consider this section as made up of the flanges only, as shown by the shaded portions 
of H, then the results are much nearer agreement, and are given in the table as jH-, 
and 2H2. Section K was also tested for two lengths of span, the results of which 
differ somewhat. If the average of the induced stresses be taken, it agrees with the 
direct tension result. 

Table II. gives the results of these tests. 



Table II. 



Beams. 


Tension Tests. 


Section. 


Area in 
sq. in. 


Vt + Vc- 


Central 
Load. 
Tons. 


Span. 
Ins. 


Fibre Stress, 

2WL 

torc=— — . 

Myt+ycH 

Tons per sq. in. 


Area in 
sq. in. 


Total 
Load. 
Tons. 


Tensile Stress 

per sq. in. 

Tons. 


A 


•7854 


•425 


•428 


36 


23-08 


•7854 


18-75 


23-87 


B 


•463 


•755 


•485 


36 


24-98 


•463 


11-62 


25-10 


C 


•805 


•805 


•805 


36 


2236 


•805 


18-8 


23-35 


D 


1-8 


•45 


•925 


36 


20-55 


•163 


3-28 


20-1 


E 


•93 


•48 


•477 


36 


1923 


•93 


23-6 


25-34 


F 


•26 


•445 


•135 


36 


21-00 


•123 


309 


25-1 


Gj 


•81 


•39 


•383 


36 


2182 


•161 


375 


23-3 


G 2 


•81 


•39 


1-055 


15 


2504 


•161 


3-75 


233 


Hi 


•448 


•266 


•109 


36 


16-43 


•108 


2-64 


24-46 


H 2 


•448 


•266 


•285 


15 


17-88 


•108 


2-64 


24-46 


iH, 


•285 


•315 


•109 


36 


21-86 


•108 


2-64 


24-46 


2^2 


•285 


•315 


•285 


15 


23-81 


•108 


2-64 


24-46 


Ki 


•67 


1-31 


•87 


36 


17-84 


•137 


2-99 


21-76 


K 2 


•67 


1-31 


2-725 


15 


23-29 


•137 


2-99 


21-76 



II. Bending Moment in Cases of Eccentric Loading, as Hooks, etc. 

Another and important case is where a section is subjected to non-axial loading, 
as in crane hooks. 

Under such conditions a section as AB of fig. 2 is generally said to experience : — 
(l) A uniformly varying stress due to the bending moment of a couple of magni- 
tude WL, the intensity at any point varying directly as the distance of 
that point from a fixed axis N in the plane of the section known as the 
neutral axis of stress ; 



216 



ME ANGUS R. FULTON ON 



(2) A direct uniformly distributed stress due to the force W acting at the centroid 
of the section AB. 

Thus if the strain produced was entirely elastic, the stress diagram of (l), when 
the material has approximately the same ultimate strengths in compression and in 
tension, would be as represented by AaN&B in fig. 3, Aa being the tensile stress 
of the extreme fibres AA, and Bb being the compressive stress of the extreme fibres 
BB of the section shown in fig. 2. 

The neutral axis NN passes through the centroid of the section. This diagram 
would be modified by the direct tensile stress AA X of (2) to the form AaiN^iB, this 



Fig 2. a 





r, gS 

having the effect of changing the neutral axis from N to N a . If NN X = x, this 
increases the external bending moment to the extent of Wx, and correspondingly 
the resisting moment of the section must be increased to the same extent. 

If t and c represent the values of the extreme stresses, then, by the ordinary 
elastic formula for determining these stresses, 

.... (2a) 



t = 



My W 
I + A 



,wp$+l 



,_My 



A V I 



I 



W 



Therefore the ratio of - = constant and the line a-fri, though its slope varies, passes 



through a fixed point N x so long as the condition of elastic stress is maintained. 



RUPTURE STRESSES IN BEAMS AND CRANE HOOKS. 217 

The very approximate nature of the results obtained by this theory of combined 
tension and bending moment is dealt with in a paper * by Mr E. S. Andrews, B.Sc., 
and Professor Karl Pearson, F.R.S., the magnitude of the error being there proved 
to depend upon the radius of curvature to which the hook is bent. They too arrive 
at the conclusion that the stress at any point in any fixed section under a given load 
is only a function of the distance of that point from the centroidal line, the maximum 
stress being obtained at the maximum distance from that line. Since the stress, then, 
depends directly on the load and this function, it follows that there will be a true 
neutral axis. An expression for its distance from the centroidal line is obtained, and 
this is a constant for a given section. The stresses, however, do not vary directly as 
the distance from this neutral line. 

If the section, which is that of a 15 -ton hook, is as shown in fig. 2a, with the 
centroid at N, then the diagram AaiNi&iB of fig. 3a represents the diagram of 
stresses to some scale, as got by Andrews' theory, right up to the true elastic limit 
on the tension side, and N x is a fixed point. 

If it were possible for the material to behave elastically right up to. the point of 
fracture, then our ultimate stress diagram in accordance with the two theories would 
be as in figs. 3 and 3a, but of greater magnitude, and would be represented by 
AaiNi&xB in figs. 4 and 4a. 

But the extreme stresses Aa x on the tensile side, as calculated from (2a) and by 
Andrews' formula and as shown on these diagrams, are much greater than the 
tensile strength of the material. Let AT and BC represent the fracture value of 
the material in direct tension and compression respectively. (While the fracture 
value in direct tension is easy to determine, that of a ductile material in direct com- 
pression cannot be determined absolutely. The term is here used to mean that value 
of the stress at which the strain exceeds a certain fraction, having regard to initial 
section and length of test piece.) Through T and C draw lines parallel to AB to 
intercept aibi in T x and Ci. Then the areas TaiT x and C&iC 1} where the stresses 
would exceed the fracture value, cannot exist. Note the area C&iCi is a minus 
quantity in fig. 4a. 

But the resisting moment of the internal stresses just before rupture must be 
equal to the external bending moment, and therefore areas within the stress limits, 
the summation of whose moments is equal to the summation of the moments of 
these excess areas, must be found to replace them. These can be represented by 
NiTiN 2 and NiCiC 2 N 2 , possibly involving another alteration of the neutral axis to N 2 . 

The conditions involved in this redistribution of stress areas are (a) that the 
summation of the moments of the complete resisting stress areas CC 2 N 2 and TT 2 N 2 
should be equal to the moment of the external load W about an axis through N 2 ; 
and (b), that the algebraic sum of the tensional and compressive stress areas should be 
equal to the external load W. 

* Drapers' Go. Research Memoirs, Technical Series I., London, 1904. 



218 



MR ANGUS E. FULTON ON 



It is now possible to proceed to establish a connection between the extreme fibre 
stresses at fracture point of a non-axially loaded body and the moment of the ex- 
ternal load. 

Let the symbols A, A t , A c , T, C, t, and c have the same meaning as in the case of 
beams, and whatever may have been the elastic distribution of stress, let the ultimate 
distribution be a rectangular one. 

If the resultant force W was very small and L very great, then only the bending 





moment stress need be considered. For this special case, if A T and A c be the tensile 
and compressive areas respectively, then 



~ = as before, for tA T - cA c = 0. 



If the ultimate stress reached be equal to the ultimate direct strength of the 
material, then 

T 



t = T ; andA x=A^ 



and A r = A 



T + C 



In order to obtain the position of the neutral axis NN under this condition, we 



RUPTURE STRESSES IN BEAMS AND CRANE HOOKS. 



219 



divide up the section of the hook in the above ratio as shown in fig. 5. Let y t and 
y c be the distances of centroids of A T and A c from the neutral axis NN. 

Then we have, since the external moment equals the internal resisting moment, 

= M TTc 2/ ' + cA fTc 2/ " 
and at the fracture point, when t and c equal T and C respectively, 



WL = *Aj^(y, + y c ) = cA_I_G/ r : „.) 



(II.) 



T + C v '" •"' T + C7 
Now, when the resultant tensile force W becomes important and is taken into 

consideration, we have 

tA t -cA a = W. 

We may imagine that this is the previous case, with L diminishing as W increases. 
Since the stress limits at fracture are T and C as before, these cannot be altered 




Fig. 5. 

to meet the resultant W, which can only be balanced by a transfer of part of the 
compression -area to the tension side. Of course, the moment of these rearranged 
areas must be equal to the external moment. Let the shaded area in fig. 5 be the part 
so transferred from compression to tension, and let the actual final position of the 
neutral axis be N 2 at a distance x from N, and z t and z c be the respective distances of the 
centroids of the areas A t and A c from those of the areas A T and A c of the special case. 
From the equation of moments we have 

W(L + x) = tk t (y t + x - z t ) + cA e (y c -x + z L ). 

Now 

C 



Aj = A T + a = A 
A f = A c - a = A 



r+c 

T 

TTc 



+ a 



y t + 



A 



and 



(». 



C 



. W(L + x) = tk^L-# t + cA^-^ + x(tA t - cA c ) - a?(t + c). 
TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 7). 



30 



220 MR ANGUS R. FULTON ON 

But 

w 

aT - a( - C) = W. . •. «(T + C) = W ; and a = ^-^ 
.'. when the ultimate stresses are T and C we have 

W(L + x) = tk^{y t + y c ) + Wx- ^L |(T + C). 

Eliminating balancing moments, this equation becomes 

CA T + C { ' J ' +y < ) = tA f% (Vt + //f) = WL + IT 
This means that what was the resisting moment of the section before this 
alteration of the neutral axis has, by reason of the rearrangement, become equal to 
the original external bending moment plus the moment of the weight about half 
the displacement of the neutral axis. > 

Since x is usually small, — - may be considered to be negligible, and therefore 

for all practical purposes the relationship between the external moment and the 
resisting moment of the section just at the point of rupture may be taken as the 
same as when there was no resultant, that is : 

WL = tA^- c (y i + y c ) = cA T ^(y t + y c ) .... (III.) 

*«T- ™i-,* + (HI.A) 

Hvt + Vc) c 

n WL T + C mT v 

corC = — — =— ..... . (lll.B) 

MVt + Vc) T 
Now, if T = C=/, as is approximately the case for W.I. and mild steel, then 

/=T = C = ^ L -._?— (3) 

Experiments. 

The results of experiments undertaken by the author with hooks made from 
various materials show that the values obtained for "/," as calculated from formula 
(3) when the ultimate stress-point is reached, agree more or less accurately with the 
direct stress values. The first tests were carried out on galvanised wrought-iron 
hooks used for suspending ship's boats. The hooks were of circular section, and had 
ends long enough to enable ordinary tension test pieces to be cut off. 

Afterwards a series of tests were made on hooks specially manufactured from 
round bars of various materials. In addition to the length necessary for the pre- 
paration of the hook, two other lengths were cut off each bar, one being tested in 
direct tension cold, and the other was first annealed before being so tested. In this 
way a direct comparison was made between the stress induced in the hook by 
bending and the ultimate tensile stress as found after heating. 

A hook was also made from a bar of rectangular section, and also another prepared 
with an approximately rectangular section by flattening the sides of a round bar. 



RUPTURE STRESSES IN BEAMS AND CRANE HOOKS. 



221 



In addition an examination was made of the investigation # by Professor Goodman 
into the strength of drop-forged steel hooks, which was published in Engineering, 
and also of the results contained in his paper f on " Crane Hooks," which followed on 
the lines of Andrews' theory and which was published in the Proceedings of the 
Institution of Civil Engineers. 

The values of (yt + y e ) in terms of the depth of section and of the consequent 
values of "f" in formula (3) is given below for the various sections investigated: — 





Description of Hook. 


(Vt + y,)- 


f. 


(1) 


Circular ...... 


■425d 


Ad 


(2) 


Rectangular ..... 


■5d 


,WL 
Ad 


(3) 


,, with rounded ends . 


•i3d 


465 A^ 


(4) 


Goodman's drop-forged steel 


1-43/i 


4-65^_ L 
Ah 


(5) 


,, wrought-iron 1st set . 


•48A 


« 7 S 

Ah 


(6) 


,, ,, 2nd set 


■i8h 


Ah 



These were found to justify the conclusions which are set forth in the present 
paper, and an analysis of these along with the experimental results are given in 
Table III. 

Table III. 















<N 


+ "1 


Max. Direct 




Description. 


Material. 


Depth. 


Area. 


Load. 


L" 




~ cr 1 

^ CO 


Tensile Stress. 


Elastic 




Ins. 


Sq. in. 


Tons. 




h3 I . <0 

te; <j & 

" 1 




Formula. 


(Cold). 


(Annealed). 


No. 1 round 


W.I. 


1-1 


•95 


4-25 


1-3 


25-97 


24-6 




46-64 


., 2 „ 


)> 


1-0 


•78 


3-90 


1-1 


25-70 


25-1 




48-58 


„ 3 „ 


B.B.H. 


1-02 


•83 


4-35 


1-0 


24-45 


23-05 


23-41 


46-61 


„ 4 „ 


Swed. 


1-02 


•82 


3-24 


1-15 


20-83 


20-00 


20-12 


39-38 


,. 5 „ 


S.M.S. 


1-00 


•78 


38 


1-1 


2520 


2500 


25-80 


46-97 


„ 6 „ 


Mild S. 


1-02 


•82 


30 


1-3 


21-92 


24-4 


24-17 


40-70 


,, 7 rect. 


1) 


•93 


•46 


25 


1-2 


27-90 


26-8 


... 


48-86 


,. 8 „ 


W.I. 


1-02 


•82 


3 

Avera 


1-3 

%e value 


21-7 


24-6 






24-20 


24-20 



* See Goodman on the " Strength of Drop-forged Crane Hooks," Engineering, lxxii. p. 537. 

t Goodman on " Maximum Stresses in Crane Hooks," Minutes of Proc. of Inst, of Civil Engineers, vol. clxvii. p. 296 



222 



MR ANGUS R. FULTON ON 



Table III. — continued. 















+ 


c-q "~ 


Max. Direct 




Description. 


Material. 


Depth. 
Ins. 


Area. 
Sq. in. 


Load. 
Tons. 


L" 


i-l 


to 
u 


Tensile Stress. 


Elastic 
Formula. 


il s 


(Cold). 


(Annealed). 


Goodman's 


Tests. 
















tons 


















^ drop-forged 


Steel 


•82 


•33 


1-76 


•95 


27-45 


no value given 


52-92 


i 




•90 


•38 


1-85 


1-04 


2604 




44-30 


i 

2 >> 




101 


•50 


2T8 


1 21 


24-18 




41-13 


1 




1-40 


•95 


7-12 


1-47 


36-60 




68-38 


H 




1-60 


1-26 


7-29 


1-83 


3069 




54-49 


2 




2-00 


1-94 


11-49 


2-10 


28-93 




52-64 


3 




2-32 


2-61 


18-15 


2-52 


35-11 




64-00 


* 




2-68 


3-34 


23-23 


2-83 


34-14 




62-27 


5 


5J 


3-00 


4-01 


26-87 
Avera< 


3-20 
*e value 


33-34 




59-86 


30-92 


5A 1st set 


W.I. 


3-49 


5-59 


325 


3-35 


23-35 


23-5 




41-6 


5B „ 


JJ 


3-47 


5-84 


34-1 


3-42 


23-93 






42-6 


10A „ 




4-36 


9-20 


57-3 


4-32 


25-4 






45-3 


10B „ 




4-34 


9-76 


57-9 


4-33 


25-4 






45-8 


15A „ 


JJ 


4-78 


12-57 


62-5 


4-65 


2014 






36-1 


15B „ 


)) 


4-97 


12-65 


900 

Averaj 


4-44 
*e value 


26-27 






47-7 


24-08 


5A 2nd set 


w.r. 


3-79 


543 


31 


3-54 


22-24 


23-4 




39-1 


5B „ 


• j 


3-36 


5-78 


33-6 


4T8 


30-16 


>) 




48-2 


10A „ 




4-75 


10-96 


54-0 


4-48 


19-37 


jj 




33-9 


10B 


a 


4-63 


10-80 


590 


4-71 


23-17 


)! 




42-1 


15A „ 


)) 


5-27 


14-53 


76-9 


5-00 


20-93 


>! 




39-9 


15B „ 


jj 


4-78 


13-51 


82-9 

Avera< 


4-84 
je value 


2591 


)! 




47-7 


23-63 



Conclusions. 

1. It may be taken as conclusive that the final distribution of stress at rupture 
point in a member subjected to an external bending moment is a rectangular one, 
unless where the cohesion of adjacent layers is not sufficient to withstand the shear 
induced by the resisting moment of the section. 

2. That, provided shear does not take place, the neutral axis moves always to 
the position which reduces the summation of the tensile and compressive stress 
areas, across a section, to the equilibrant of the external forces. (In the case of 



RUPTURE STRESSES IN BEAMS AND CRANE HOOKS. 



223 



a beam this reduces to zero ; in that of a hook, at the principal section to the 
suspended weight.) 

3. That the total resisting moment of these stresses must be equal to the ex- 
ternal bending moment as measured to the neutral axis at rupture point, but that 
these balancing moments do not differ materially from those measured to an axis 
obtained by dividing the sectional area into tensile and compressive stress areas 
which are in inverse proportion to the magnitude of their respective ultimate direct 
stresses. 

The advantage of these formulae are important. It is possible to indicate with 
certainty the magnitude of the load which will cause rupture in a beam or a hook 
provided there is known the point of application or the effective arm of the load, 
the cross-section of the beam or hook, and the breaking strengths of the material 
when subjected to the different forms of direct loading. 



TRANS. ROY SOC. EDIN., VOL. L. PART I. (NO. 7). 



31 



( 225 ) 



VIII. — Scottish National Antarctic Expedition : A Description of the Systematic 
Anatomy of a Foetal Sea- Leopard (Stenorhynchus leptonyx), with Remarks upon 
the Microscopical Anatomy of some of the Organs. By Harold Axel Haig, 
M.B., B.S., M.R.C.S., late Lecturer in Histology and Embryology, University 
College, Cardiff; M'Robert Research Fellow, University of Aberdeen. Com- 
municated by Professor Arthur Robinson, M.D. 

(MS. received January 26, 1914. Read February 16, 1914. Issued separately April 30, 1914.) 

[Plates XIX.-XXIL] 

During the Scottish Antarctic Expedition of 1892-93 Dr W. S. Bruce secured 
foetuses of Stenorhynchus leptonyx and Lobodon Carcinophaga, and on his return 
passed them over, with other material, to Professor DArcy W. Thompson for the 
Zoological Museum of University College, Dundee. While some of the material has 
unfortunately been lost sight of during these twenty-one years, one specimen, viz. that 
of a foetus of Stenorhynchus leptonyx, was still in existence, and was returned by 
Professor D'Arcy Thompson to Dr Bruce, who in turn asked me to examine and 
report upon it. Furthermore, during the voyage of the Scotia embryos of Leptony- 
chotes weddelli were obtained by the Scotia naturalists, and these were passed on for 
description to Professor Waterston, at that time in the University of Edinburgh. It 
is on this material that the present monograph is based. 

Preliminary Considerations. 

The foetus of Stenoryhnchus leptonyx, which is in a good state of preservation, 
measures 122 mm. from the tip of the tail to the most prominent part of the mid- 
brain ; the greatest breadth is about 43 mm., and the greatest dorsi- ventral measure- 
ment is in the mid-dorsal region, measurino; 35 mm. The umbilicus is situated 36 mm. 
from the cloacal aperture, the umbilical cord being relatively short, owing to the 
fact that the umbilical vessels soon undergo division into several large branches ; a 
portion of the placenta with fragments of the foetal membranes is still attached 
to the cord. The actual mode of placentation and the disposition of the foetal 
membranes are points which will be discussed at a later stage (see infra, on the 
Placenta). 

The skin is in many places thrown into folds, some of these being normal, but 

others undoubtedly due to shrinkage consequent upon the action of the preservative 

medium. The nippers are fully formed, nail rudiments being present upon the 

dorsal aspects of the distal phalanges. 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 8). 32 



226 MR HAROLD AXEL HAIG ON 

The head is at this stage not large in proportion to the trunk ; rudiments 
of the vibrissse are to be seen at the sides of the snout, and the eyelids are 
formed although the palpebral fissure is not as yet open. The tongue is an 
elongated organ, with a cleft tip, the two divisions appearing between the lips in 
the middle line. 

No sign of an external ear is present, nor is there any opening indicating the 
position of an auditory meatus. 

The whole trunk is curved towards the ventral aspect, but some of the curvature 
is probably due to mechanical causes subsequent to removal of the foetus from 
the uterus. 

Reference to PI. XIX. fig. 1 will render the above points clear. 

SECTION I. 

General Topography and Anatomy. 
A. Appearance of the Main Viscera in situ, from the Ventral Aspect. 

A median ventral incision was made, and flaps of skin and deeper tissues turned 
back to expose the structures in the neck, thorax, and abdomen ; the sternum and 
ribs being also removed for the purpose of demonstrating the thoracic viscera, whilst 
the attachments of the diaphragm to the lower ribs were likewise severed. A 
separate flap was raised in the neck region to expose the larynx, trachea, thyroid, and 
parathyroids, and finally the parietal pericardium was cut away. 

In the region of the umbilicus, care was taken not to sever the connection of the 
urinary bladder with the umbilical cord, and the umbilical vein passing from the 
cord to the liver was also kept intact ; subsequently, however, these connections were 
severed for greater convenience of examination. 

Reference to fig. 2, Plate XIX., will show that the heart is at this stage a large 
organ filling the greater part of the thoracic cavity ; from the ventral aspect, the 
right ventricle appears larger than the left, and the right auricular appendix wider 
than the left ; the right appendix has a deep notch in its lower border, whilst the 
left one possesses three such notches. 

A portion of the aortic trunk shows above and dorsal to the right appendix. 

The thymus is relatively large, and extends from the root of the neck, where it is 
bifurcated, towards the left of the middle line, until it reaches a point just anterior 
to the left auricular appendix : the main mass of the thymus is subdivided into a 
number of lobes and lobules, and there are a few small isolated masses situated at 
the sides of the trachea just anterior to the upper bifurcated extremity.* 

The lungs lie compressed against the walls of the thorax, and are not very obvious 
from the ventral aspect, the right upper lobe being most prominent, and below 

* Microscopical examination showed that these isolated masses possessed a typical thymus structure. 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 227 

this a small portion of the middle lobe, whilst none of the lower lobe is visible ; 
both lobes of the left lung show, the anterior edges and portions of the lateral 
surfaces being seen. In the neck region the larynx and trachea form prominent 
features, whilst at the sides of the trachea the lateral lobes of the thyroid gland 
with the lower parathyroids are to be seen ; no thyroid isthmus is, however, to be 
detected, a point which is noteworthy. 

The abdominal viscera from the ventral aspect (PL XIX. fig. 2) : — The liver 
forms the most prominent organ in the abdomen, its right and left lobes, together 
with certain accessory lobes, occupying about one-half the available space ; the right 
lobe appears to be larger than the left, but in reality this is not the case, since, when 
viewed from the dorsal aspect (see PI. XXI. fig. 5), the left lobe is seen to be much 
the bulkier of the two. A fissure passes obliquely downwards and inwards from the 
middle of the lateral aspect of the right lobe, and effects a partial subdivision of this 
lobe into two, but the cleft does not extend deeply into its substance ; whilst a 
small flap of the upper of the two subdivisions is seen a short distance internal to 
the right lateral margin, and partly conceals an aperture in the lobe in which the 
fundus of the gall-bladder appears (PI. XIX. fig. 2 15 ). 

Between the right and left lobes there is a fairly wide cleft, in which may be seen the 
umbilical vein passing from the umbilicus towards the ductus venosus. 

Coils of small intestine are seen lying caudal to the liver, but the stomach is not 
visible, being largely hidden by the left hepatic lobe. 

The urinary bladder is a very elongated structure, and is attached ventrally to 
the umbilicus : it opens caudally into the cloaca. The umbilical arteries pass along 
the lateral aspects of the viscus towards their destination in the placenta, and are 
well seen in transverse sections. (See PI. XXI. fig. 4.) 

The chief points of interest in connection with the above description of the 
ventral aspect of the viscera are : — 

(i.) The relative size of the thymus, which, although as a rule large at this stage 
of development, appears in the present case to be markedly so ; the left 
lateral deviation of its caudal extremity is also a point worthy of note. 
The presence of isolated lobules of this gland suggests a possible origin of 
these from some of the higher branchial clefts. 

(ii.) The deep notching of the auricular appendices in the heart is a feature which 
seems very striking upon opening the pericardial cavity. One other point 
also, viz. the well-marked interventricular furrow, is a characteristic which 
becomes more marked as development proceeds, the external subdivision of 
the ventricular portion of the heart giving it in the full-grown seal an 
almost bifid appearance. 

(iii.) The great longitudinal extent of the liver, an organ occupying at this stage of 
development a relatively large proportion of the abdominal cavity. 



228 MR HAROLD AXEL HAIG ON 

(iv.) The peculiar shape of the urinary bladder. According to Hepburn,* the 
bladder in the adult seal is represented by the whole length of the foetal 
organ, since no shrinkage takes place in the cephalic portion to form a 
definite urachus. 

B. General Topographical Anatomy of the Head. 

I. The oral, nasal, and cranial cavities, with their contents (PL XX.). 

A median sagittal section of the head was made, so that the oral, nasal, and 
cranial cavities were laid bare ; whilst, in order to expose the contents of the nasal 
fossae, the nasal septum was subsequently removed. 

Rudiments of the incisor milk-teeth were cut through in the upper and lower 
jaws, and the surface of the mesial section of the hypophysis cerebri exposed, the 
hypophysis lying in the sella turcica of the developing sphenoid bone at the base of 
the skull (PL XX. fig. 3 3 ). 

Other points worthy of note in such a hemisection of the head are : — 

(a) The elongated tongue (PL XX. fig. 3 2 ). 

(b) The falx cerebri, covering the mesial surface of the right cerebral hemisphere 

(PL XX. fig. 3 9 ) ; about the middle of the concave edge of the falx the 
corpus callosum is seen in section (PL XX. fig. 3 9/ .) 

(c) The optic thalamus (PL XX. fig. 3 8 ) continued posteriorly into the 

mesencephalon and pons ; dorsal to these latter, the Sylvian aqueduct with 
its roof, in which the rudiments of the corpora quadrigemina (fig. 3 7 ) 
are to be made out ; posterior to these are seen the mesial section of the 
cerebellar vermis, and, ventral to this, the 4th ventricle and medulla 
oblongata (PL XX. fig. 3 3 - 6 ' 6 ' 4 ). 

(d) The cavity of the 3rd ventricle, with its extension into the infundibulum of 

the hypophysis. 

After the removal of the falx cerebri and nasal septum, the following additional 
structures come into view : — 

In the cranial cavity (PL XX. fig. 5) : — 

(e) The mesial surface of the cerebral hemisphere (h.). 

(f) The olfactory lobe (olf. I.), lying between the fore-brain and the anterior 

boundary of the cranial cavity : the lobe is not a large one, and does not 
appear to give off many nerve-filaments to the ethmoidal region of the 
nasal fossa. 

In the region of the nasal fossae : — 

(g) The rudiment of the ethmo-turbinal bone (eth. t.) lying at the superior angle; 

this rudiment, which at the present stage is cartilaginous, shows three ridges 

* Tunis. Iltxj. Hoc. EiHn n vol. xlviii. part i., No. 3, 1913. 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 229 

with two intervening depressions, and represents the olfactory region of the 
nasal fossa. 
(h) The large maxillo-turbinal rudiment (mx. t.), presenting upon its mesial 
surface a number of narrow lamellae with intervening sulci : it is carti- 
laginous and is covered by a mucous membrane lined by ciliated epithe- 
lium. Anteriorly the maxillo-turbinal bone is attached to the outer wall 
of the fossa, whilst posteriorly it fuses by an elongated pedicle with the 
periosteum of the base of the skull. 

The buccal cavity is lined by a mucous membrane covered by stratified epithelium 
with but few cell-layers. At the present stage of development, the membrane bones 
of the cranial vault are partially ossified, but the bones of the base of the skull (basi- 
sphenoid, basi-occipital) are still partly cartilaginous ; the sphenoid bone (see text-fig. 
3, b) is in the " irruption "-stage of endochondral ossification, the process being seem- 
ingly delayed by the rather late persistence of the epithelial connection between the 
hypophysis cerebri and the buccal epithelium.* 

The brain. — General superficial anatomy (PI. XX. figs. 1 and 2). The brain of 
the present specimen shows some very well-defined characters, and on the whole 
may be said to conform to the mammalian type : viewed from the dorsal aspect 
(PI. XX. fig. l), the following features are obvious : — 

(i.) The large cerebral hemispheres separated by the longitudinal sulcus in which 

the falx cerebri is normally lodged, 
(ii.) Behind the hemispheres, the crura cerebri and region of the mesencephalon, 

covered by fragments of torn pia mater, 
(iii.) The cerebellum, consisting of the mesially situated rudiment of the vermis, 

and on either side of this the cerebellar hemisphere : the latter already 

present a number of flattened laminae with intervening sulci, 
(iv.) The 4th ventricle in its lower half, with the restiform bodies bounding it 

on either side ; the floor of the ventricle is seen owing to previous removal 

of the roof of pia mater. 

The surface of each cerebral hemisphere is quite smooth upon its superior aspect, 
and shows no indications as yet of any convolutions. 

Viewed from the side, several additional features become apparent, viz. (PI. XX. 
fig. 2) :- 

(v.) The olfactory lobe, and its connection with the ventral part of the fore-brain, 
(vi.) A wide shallow depression passing from the lower aspect of the fore-brain 
upwards and backwards towards the hind-brain (future occipital lobe) : 
this depression indicates the position of the future Sylvian fissure (PL XX. 
fig. 2 2 ), and ventral to its posterior extremity a forward extension of the 
hemisphere represents the rudimentary temporo-sphenoidal lobe. 

* See also P. T. Herring on " Development of Mammalian Pituitary Body," Joum. Exper. Physiol., 1909. 



230 MR HAROLD AXEL HAIG ON 

Upon the ventral aspect, at the base of the brain, the infundibulum forms a down- 
ward projection from the floor of the 3rd ventricle (PI. XX. fig. 2 3 ), whilst half way 
between the infundibulum and a projection which marks the position of the ventral 
part of the pons the 3rd nerve forms a noticeable feature (PI. XX. fig. 2 4 ). 

Additional features to be made out in a sagittal section taken through a cerebral 
hemisphere, the optic thalamus and mid- and hind-brain (PL XX. fig. 4) : — In such 
a section the general relations of the various regions of the brain may be studied to 
a certain extent, and, moreover, by staining with hsematoxylin and eosin the main 
distribution of grey and white matter at this stage of development may be made out. 
The cortex cerebri (PI. XX. fig. 4 4 ' °) is made up of two parts, viz. : — 

(i.) A superficial layer of white fibres and neuroglia, forming a very narrow zone 

(PI. XX. fig. 4 5 ). 
(ii.) A deeper layer of deeply-staining nerve-cells, with neuroglia, of somewhat 

wider extent than (i.). 

Deep to the cortex comes the white matter of the hemisphere, which is at this 
stage not very thick. A section taken so as to pass completely through the cavity of 
the prosencephalon shows in addition an inner layer, somewhat deeply stained, 
which later on will form the ependymal lining of the lateral ventricle. 

Below the main mass of the hemisphere, part of the descending horn of the lateral 
ventricle is seen (PI. XX. fig. 4"), and this becomes continuous anteriorly with 
the rhinencephalon, passing into the olfactory lobe. The anterior boundary of the 
descending horn is formed by a layer of grey matter, which above takes the form of 
radiating streaks, alternating with white matter. 

In the region of the optic thalamus, small isolated patches of grey substance 
appear near the dorsal surface, but in the region of the corpora quadrigemina white 
matter seems as yet to predominate. 

The grey matter of the cerebellar hemisphere appears to lie chiefly on the surface, 
but there is a deeper zone of small nerve-cells, the two layers being separated by a 
clear zone of white matter : the cells of Purkinje do not as yet appear to have become 
differentiated as a distinct line of neuroblasts. In the region of the medulla oblongata, 
the nuclei of the 10th and 12th cranial nerves form a series of groups of rather large 
nerve-cells (PI. XX. fig. 4 1 ) ; whilst the pyramidal tract appears as a well-defined 
longitudinal set of fibres. 

The infundibulum, with a small portion of the cavity of the 3rd ventricle, has been 
already noted ; the ventral mass of the pons is a very obvious feature lying just 
anterior to the medulla (PI. XX. fig. 4). 

A fold of pia mater (PI. XX. fig. 4 3 ) shows between the overhanging posterior 
extremity of the cerebral hemisphere and the mesencephalon, whilst in the deep 
fissure anterior to the optic thalamus a small piece of choroid plexus appears. 

Points for comparison with the brains of other carnivora are the following : — 



THE SYSTEMATIC ANATOMY OF A F03TAL SEA-LEOPARD. 231 

(a) The relative shortness of the corpus callosum, a structure which, for instance, in 

the cat, at a corresponding stage of development has assumed a much longer 
antero-posterior measurement ; moreover, it would seem that in this Seal 
embryo the corpus callosum is taking a more vertical course than is usual. 

(b) Altogether the general appearance and stage of development of the brain under 

discussion corresponds very closely with one figured by His * of a three- 
months human foetus, with the exceptions that the hypophysis cerebri is 
much further advanced (see infra) and the cerebellum has assumed greater 
complexity. 

C. Anatomical Details of the Remaining Viscera. 

The thyroid and parathyroids, the thymus, pancreas, and tongue, will be described 

under the histological section of this pamphlet ; in the present instance the following 

organs will be considered, and comparisons made with other types, where this is 

possible : — 

(i.) The lungs. 

(ii.) The heart. 

(iii.) The liver, stomach, and intestines. 
(iv.) The spleen. 

(v.) The kidneys and adrenal bodies, 
(vi.) The genital glands and ducts : the urinary bladder. 

The histology of some of these will be dealt with later. 

(i.) The lungs (PI. XXI. fig. 3). — -Both lungs are somewhat compressed against 
the thoracic walls, owing to the large size of the heart, and it is only upon removing 
the latter organ that a good view can be obtained of the roots of the lungs and their 
ventral aspect ; posteriorly the inner margins of the upper and lower lobes of both 
lungs are grooved longitudinally by the vertebral column, whilst laterally the surfaces 
of all the lobes are grooved by the ribs. 

In the right lung, three lobes, upper, middle, and lower, can be distinguished f ; 
the middle lobe is peculiar in that at the root it gives off two caudally directed 
subdivisions, the largest of which is almost a separate lobe, and presents dorsal, 
right lateral, and anterior surfaces (PI. XXI. fig. 3 a). The cranial and caudal lobes 
of both lungs are somewhat similar in shape, the caudal lobes being the larger, being 
not unlike the corresponding lobes of a human lung. Marked trabecule of con- 
nective tissue are to be seen upon the surfaces of all lobes, with finer strands passing 
off in all directions, and these subdivide the surface into polygonal areas, the bases, 
as it were, of the superficial lobules ; on being placed in water the lungs sink. 

At the root of the lung the bronchi and pulmonary vessels are seen entering (or 

* Hertwig, Handbuch der JEntwickelungslehre der Wirbeltiere. 

t The terms " upper," etc., are here used with reference to the vertical position of the trunk, and not the natural 
position of the adult animal. 



232 MR HAROLD AXEL HAIG ON 

emerging from) the upper lobes, the bronchi being dorsal to the pulmonary arteries, 
although the left bronchus is much closer to the artery than the left. 

(ii.) The heart (PL XX. fig. 2 and text-fig. l). — All four cavities of the heart 
contain firm clot, which extends into the auricular appendices ; on section in the 
coronal plane, the cavities of the auricles appear to be larger than those of the 
ventricles, whilst the auricular appendices add considerably to the auricular capacity. 
Moreover, although from the ventral aspect the right ventricle appear larger than 
the left, there is not the same relative difference between the capacities of the 
ventricles, both appearing to possess much the same size in median coronal section ; 
the thickness of the myocardium is rather greater in the left than in the right 
ventricle, that of the auricles being about equal on both sides. The aortic bulb is 



Fig. 1.— Dissection to show the relations of the aortic trunk, pulmonary trunk, and ductus arteriosus. 



R.S. Right subclavian. 
AO. Aorta. 

P. A. Pulmonary artery. 
D.A. Ductus arteriosus. 



L.V. Left ventricle in section. 
R.V. Right ventricle in section. 
B.AO. Aortic trunk and bulb. 
IN. Innominate trunk. 
R.C. Right common carotid. 

The dotted lines show the connection of the pulmonary trunk with the right ventricle ; 
the lumen of the aortic bulb is opened, and two semilunar valves are seen. 

full of firm clot, and passes at once into a relatively short but thick aortic arch ; the 
pulmonary trunk shows the same relations to the aortic trunk as it does in the case 
of the human foetus of a corresponding stage of development, whilst the ductus 
arteriosus is relatively wide and is a continuation of the main pulmonary trunk after 
the latter has given off the two branches to the lungs (see fig. 2). The ductus joins 
the arch of the aorta close to the origin of the latter from the bulbus aortse. 

The cardiac valves are well developed, the semilunar valves consisting of thin 
plates of fibrous tissue, the free edges of which project into the aortic and pulmonary 
trunks, whilst the mitral and tricuspid valves have their free edges projecting into 
the ventricular cavities. The columnse carnse are not marked features, nor do the 
chordae tendinse appear to be strongly developed at this stage. 



THE SYSTEMATIC ANATOMY OF A FXETAL SEA-LEOPARD. 233 

Serial sections taken coronally through the heart show that the foramen ovale is 
a feature at this phase, but the aperture is a very narrow one, and will not admit a 
glass seeker of more than 1 mm. diameter. 

From the above description it will be seen that the heart of Stenorhynchus during 
early foetal life corresponds more or less closely with the typical mammalian organ 
in its developmental aspects ; the interventricular furrow, however, is a marked ex- 
ternal feature, and during development, at least in the Weddell Seal, it becomes 
deeper, so that the full-grown heart possesses a bifid apex, the tips of the two 
ventricles being separated by a deep notch. # 

(iii.) The liver, stomach, and intestines (PI. XXI. figs. 1 and 5). — In some respects 
the liver of this specimen shows characters similar to those of the human fcetal organ, 
but the accessory lobes, and the clefts which produce these, are to a certain extent 
atypical ; thus the right lobe shows from the ventral aspect a fissure which passes 
obliquely inwards from the lateral surface, but this fissure is not seen from behind, 
as it extends for only a short distance into the substance of the lobe. 

From the dorsal aspect (PI. XXI. fig. 5) it appears that the left lobe is the bulkier 
of the two, and the inner surface of this lobe shows depressions corresponding to 
the ventral surface of the stomach, a small area of the spleen, and anteriorly the 
oesophagus and vena cava ; the inner surface of the right lobe shows dorsally de- 
pressions corresponding to the numerous subdivisions of the right kidney, whilst the 
lateral aspects of both lobes are grooved by five or six of the posterior ribs. In the 
middle line ventrally the umbilical vein (PI. XXL fig. 5, l.v.) forms a thick cord, 
passing towards the ductus venosus. 

The gall-bladder is not seen from the dorsal aspect except when the lobes are 
widely separated, when it appears as an elongated sac, deeply embedded in the deep 
surface of the right lobe, the fundus presenting ventrally in the small opening noted 
in the topographical description (see supra). 

Certain accessory lobes show up when the two main lobes are separated from one 
another : these may possibly be the homologues of the Spigelian and quadrate lobes 
of the human organ, but their relations are somewhat different. 
The main points for comparison to be noted in this organ are : — 

(i.) The relative longitudinal extent of both lobes, this being distinctly greater 

than is the case with most other carnivora. 
(ii.) The small accessory flap guarding the aperture in which, ventrally, the fundus 

of the gall-bladder is to be seen, 
(iii.) The large volume of the abdominal cavity, occupied by the whole liver, at a 
stage when other abdominal viscera have assumed a relative importance 
in size. 

The stomach and intestines (PI. XXI. fig. l) are seen from the ventral aspect, after 

* See Hepburn, Trans. Roy. Soc. Edin., vol. xlviii., 1913. 
TRANS. ROY. SOC. EDIN, VOL. L. PART I. (NO. 8). 33 



234 MR HAROLD AXEL HAIG ON 

removal of the liver, by which organ they are largely concealed ; the stomach is 
placed with its long axis nearly parallel to the long axis of the foetus, and possesses 
a well-marked cardiac extremity lying with its fundus pressed against the diaphragm, 
and a pyloric end which is narrow and passes by a sharply curved portion into the 
first part of the small intestine. The lesser curvature of the stomach looks towards 
the ventral aspect, the greater curvature and fundus are dorsally situated, whilst 
a peritoneal fold, the representative of a great omentum, is attached to the whole 
length of the larger curvature ; the lesser curvature and the duodenum have 
passing between them a narrow peritoneal sheet, which holds up the duodenum so 
that its first part runs parallel with the stomach. 

The stomach is an inch and a half in length, and at its cardiac end is joined by 
the oesophagus, the latter being a somewhat wide tube, two inches long and quarter 
of an inch in diameter. 

The first part of the small intestine (duodenum) makes three bends, and possesses 
four distinct portions, the first of which is parallel with the smaller curvature of the 
stomach, whilst the second, third, and fourth divisions enclose a portion of peritoneum, 
between the layers of which the pancreas is held in position. The coils of small 
intestine proper are already many in number, and their general arrangement may be 
made out by reference to PL XXI. fig. 1. 

With regard to the large intestine, the position of the caecal pouch is noteworthy : 
this pouch is placed opposite the level of the third bend of the duodenum, being con- 
nected with the latter by a short fold of peritoneum. No sign of a vermiform ap- 
pendix is to be made out ; as a matter of fact, this organ is not seen in the full-grown 
animal.* 

(iv.) The spleen (PI. XXI. fig. 6) has a situation upon the left side of the abdomen, 
parallel to the greater curvature of the stomach and attached to this by a fold of 
peritoneum ; in thickness this organ does not measure more than one-eighth of an inch, 
but in length exceeds two inches. There are no notches in either its ventral or its 
dorsal edge, and the hilus occupies a large proportion of a ridge forming its inner 
margin which lies close to the stomach ; the outer surface of the spleen is grooved 
by two or three of the posterior ribs. 

(v.) The kidneys (PI. XXI. fig. 4). — These organs are somewhat elongated oval 
bodies lying low down at the back of the abdominal cavity and close to the middle 
line ; each kidney belongs to the type common to the Pinnipedia, viz. the per- 
manently subdivided type, where the organ is made up of a large number of anatomi- 
cally distinct renal pyramids, t the secreting tubules of which open into a common 
pelvis, from which latter a ureter conducts the secretion to the urinary bladder. 

In this specimen there are in each kidney about two hundred and forty small 

* Hepburn (Trims. Roy. Soc. Edin., vol. xlviii. part i., No. 3, 1913) regards the csecal diverticulum as a combined 
caecum and vermiform appendix, 
t " Renculi " of German authors. 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 235 

raised areas, circular in contour, representing the bases of the renal pyramids : these 
are mostly uniform in size, but some, smaller than the majority, lie rather below the 
general surface. A mesial coronal section of the kidney passes through many of 
the pyramids and also opens up the calyces and pelvis ; according to Chievitz,* a 
certain amount of reduction takes place during development, so that whereas in certain 
instances about two hundred calyces may be present, in the full-grown animal only 
one hundred and forty remain, the reduction apparently commencing in those of the 
6th and 7th order of origin. 

The ureter emerges from about the middle of the inner and dorsal margin of the 
kidney, and passes down parallel to the mid-line, crossing the Miillerian ducts dorsally at 
right angles, and finally opens dorsally into the lower segment of the bladder ; the 
hilus of the kidney from which the ureter emerges and into which the renal vessels 
pass is not a marked feature. The "pelvis" mentioned above is also but little 
developed, since the ureter divides almost at once into two main branches, the latter 
undergoing further subdivision in the kidney until the final divisions are reached 
close to the surface of the organ in the cortical zone of each renal pyramid. 
Each kidney measures about one inch by half an inch. The adrenal bodies (PI. XXI. 
fig. 4) are small reniform structures lying one on each side just anterior to the 
kidney ; there is a loose connection with the latter organ by means of a peritoneal 
band. In actual shape the adrenal is a flattened pyramid with a convex base facing 
outwards and ventralwards, whilst the hilus is placed at about the middle of the 
inner, or rather dorsal, edge, close to the vertebral column ; the length of each gland 
is half an inch, and the breadth one quarter inch. Some points in the histology of 
both the kidney and the adrenal gland will be described later. 

(vi.) The genital glands with their ducts (PI. XXI. fig. 4) are in the present 
specimen already sufficiently established to be able to determine the sex of the animal 
— i.e. they are ovaries, and the Fallopian tubes pass from the outer ends of the glands 
to fuse in the middle line dorsal to the bladder and ventral to the rectum, and from 
the rudimentary uterus ; the outer ends of the Fallopian tubes are dilated to form 
large ampullae for the reception of the ova, and these ampullae lie ventral to the 
outer pole of each ovary. 

Each gland is an ovoid body lying obliquely from without backwards and inwards 
just behind the posterior pole of the kidney ; from its anterior extremity an elongated 
narrow muscular band, the diaphragmatic ligament of the mesonephros, passes 
towards the diaphragm and becomes attached to the dorsal wall of the abdomen. 

The urinary bladder (PI. XXI. fig. 4 3 ) is an elongated organ attached above by a 
patent allantoic duct to the umbilicus ; on transverse section, three apertures are seen 
— a median one, the bladder, and two lateral, the lumina of the umbilical arteries. 
The latter are passing to the umbilicus from their origin in the aorta. 

The bladder opens into the cloaca by a transversely elongated slit-like aperture, 

* Archiv Anat. u. Embryol., Supplement, 1897. 



236 MR HAROLD AXEL HAIG ON 

and the ureters are to be seen opening into the bladder laterally upon its dorsal 
aspect ; it is noteworthy that in the seals the bladder is represented by the entire 
intra-abdominal extent of the allantois.* 

Summary of the Anatomical Features. 

From the foregoing description it will be readily gathered that the Sea-Leopard 
Seal, during the middle phase of foetal life, presents fairly typical embryological 
features ; the age of the present specimen can hardly be worked out with any 
approach to accuracy, but it may be stated that the stage of development of most 
of the organs would place the foetus at about the end of the first third of intra- 
uterine life. The exact period of gestation of seals is, however, somewhat difficult 
to determine owing to the peculiar habits of mating which these animals have, so 
that the above estimate should be accepted with some reservation. 

In summarising the main anatomical features, it is possible to pick out the 
following more obvious characters : — 

(a) In the brain : firstly, the relatively advanced stage of the cerebellum, the 

hemispheres of this portion showing distinct evidence of folds and sulci 
which are not to be made out in a three-months human foetus ; secondly, 
the comparatively advanced development of the pituitary body, a feature 
which will be more readily appreciated when the histology of the hypo- 
physis is considered (see infra). 

(b) In the heart : the most prominent feature is the late persistence of the bulbus 

aortse, whilst the peculiar shape of the auricular appendices is also worthy 
of note. 

(c) In the lungs : the possession of an accessory lobe by the right lung, and the 

relative size of these organs, which are certainly large, are points of com- 
parative value. 

(d) Other points which may be emphasised are the large size of the thymus and 

the left-sided deviation of this organ, the apparent absence of a thyroid 
isthmus, and the large size of the lowest parathyroids ; in connection with 
the alimentary tract, the forward position of the csecal diverticulum and 
the great length of the lobes of the liver. The kidneys are noteworthy on 
account of their surface lobulation into numerous renal pyramids, whilst 
the adrenal bodies do not lie directly upon the anterior poles of the kidneys. 
Moreover, the adrenals are relatively much smaller than is. the case with 
these bodies in the human foetus at a corresponding stage of development. 
Lastly, certain external characters, such as the absence of an external ear and 
the protruding bifid tip of the tongue, are features so obvious as to need 
no further comment. 

* Hkfbukw, Trans. Roy. Soc. Edin., vol. xlviii. part i., No. 3, 1913. 



THE SYSTEMATIC ANATOMY OF A F03TAL SEA-LEOPARD. 237 

SECTION II. 

An Outline of the Microscopical Anatomy of some of the Organs. 

The histological characters of certain of the viscera were examined in this foetus, 
and in a few instances, viz. kidney and pituitary body, were found interesting from 
the point of view of the histogenesis of their essential secreting portions. The 
following account must, however, be looked upon as purely descriptive in character, 
since from the mere observation of features presented by a single specimen at one 
stage of development it is hardly possible to formulate a complete account of the 
histogenesis of any one organ. 

(i.) The thyroid and 'parathyroids (see PI. XXI. fig. 2). — The lateral lobes of the 
thyroid gland are situated rather far forward in the neck region, reaching the level 
of the lower border of the cricoid cartilage ; each lobe measures about half an inch 
in length. The largest parathyroid is the posterior one, and is placed upon the 
mesial surface of the thyroid at the ventral and posterior angle of the lobe. The 
anterior parathyroid is quite small, and is found about the middle of the dorsal 
margin of the lateral lobe. 

In minute structure the thyroid is seen to be made up of a large number of small 
vesicles, some of them showing evidence of recent origin from branched tubular 
columns of cells, and in some of them colloid is already to be detected ; the vesicles 
are lined by a cubical epithelium, and there is a small amount of interstitial connec- 
tive tissue, but no sign of a basement membrane outside the epithelium. The whole 
lobe is subdivided into relatively few rather large lobules by coarser trabeculse of 
connective tissue which are given off from the inner surface of a rudimentary 
capsule which surrounds the lobes. Blood-vessels are fairly numerous, the larger 
branches being supported by the coarser trabeculse of connective tissue lying 
between the lobules. 

The parathyroid (PI. XXI. fig. 2, p.th.) shows a quite typical structure : it is 
enclosed by a capsule of somewhat open connective tissue, which gives off trabeculse 
passing into the interior of the gland. The essential secreting cells occur in the form 
of folded columns of cuboidal or polyhedral cells, whilst these columns are in close 
apposition to the blood-channels, the latter being at this stage lined by a definite 
endothelium ; the latter, however, is not so marked a feature as it is in the blood- 
channels occurring in the pars anterior of the pituitary body of this foetus. 

(ii.) The thymus shows on section and microscopical examination a number of 
lobulated masses of lymphoid tissue supported by a framework of open connective 
tissue in which run a few rather large blood-vessels ; here and there in the actual 
substance of the lymphoid masses there occur a few areas which appear clearer and 
which probably represent the remains 1 of the original lumina of the epithelial tubules 
from which the gland arises. 



238 MR HAROLD AXEL HAIG ON 

The accessory portions of the thymus, noted in the general description of the 
viscera as being isolated from the main mass, are in all probability derived from the 
2nd branchial cleft, and do not fuse with the portions derived from the 3rd and 4th 
clefts. 

Groschuff and Verdun state that the thymus in carnivora arises invariably from 
the 3rd and 4th clefts, but in the rabbit, according to Verdun, additional parts may 
arise from the 2nd cleft.* In the present instance, paired accessory portions were 
found lying dorsal to the sterno-mastoid muscle on either side of the trachea, and 
these on examination showed a typical thymus structure. 

The lymphoid nodules of the thymus present a more or less uniform density with 
the exception of the occurrence of the above-mentioned clearer areas, and no sub- 
division into cortex and medulla is as yet obvious ; nor are any corpuscles of 
Hassall to be observed. 

(iii.) The lungs (PI. XXII. fig. 3). — Microscopical examination reveals in these 
organs a structure entirely comparable to that of a compound tubular gland, the 
branching tubules of which lie embedded in connective tissue ; the latter exists in 
large amount and is of a fibro-cellular character. The epithelium lining all of the 
ramifications of the bronchi is of the high columnar type, with the nuclei lying next 
the basement membrane, whilst immediately outside the latter there is seen a fairly 
wide zone of tissue, more densely cellular than the true interstitial connective tissue ; 
this denser zone is the anlage of the fibro-muscular and elastic coats of the bronchioles. 

The epithelium of the branching tubules is ciliated, and it is only during later 
stages that the cilia disappear in those portions of the bronchioles where the latter 
expand into the infundibula and alveoli ; at the present stage, although in some places 
the tubules appear to widen out into sac-like expansions, the epithelium remains of 
the ciliated variety, since no true alveoli with air-sacs are as yet developed. 
The interstitial connective tissue, which during later stages becomes compressed 
and relatively diminished in amount by the preponderating development of the 
alveoli, contains some large blood-vessels, but these are as yet relatively few in 
number. 

(iv.) The pancreas (PI. XXII. fig. 5). — The histological features presented by this 
gland are quite typical : branching tubules supported by a fine meshwork of con- 
nective tissue, the whole enclosed by a capsule of somewhat open character, from 
which trabeculse pass into the substance of the gland. The tubules are lined by 
columnar epithelium, but there is as yet no definite basement membrane ; t the 
lumina of the developing alveoli are quite small. The blood-vessels are quite small 
and apparently not very numerous at this stage ; no signs of any cell-groups com- 
parable to islets of Langerhans are to be detected, but probably it is too early for 

* See Hertwig, Handhuch der Entwichelungslehre der Wirbeltiere, 1906. 

t The. connective tissue appeared to have shrunken away from the tubules, leaving a considerable space between the 
two (see fig. 5). 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 239 

these to have been formed. When a section of the pancreas stained with hsema- 
toxylin is examined, it appears that the cells lining the alveoli are not characterised 
by the deeply-staining outer zone which is so marked a feature in the pancreas 
of the armadillo ; moreover, the nuclei lie at the outer or attached borders of 
the cells. 

(v.) The spleen. — In minute structure, the spleen presents the following features : — 

(a) Externally, a somewhat dense fibrous capsule, in which also unstriped muscle 

cells are undergoing development. 

(b) Trabeculse passing from the capsule into the substance of the gland, and 

forming a network in the interior : near the surface some large arteries 
may be seen passing in, more especially in the vicinity of the hilus, and 
corresponding venous branches are emerging. 

(c) The bulk of the organ is made up of a mass of erythrocytes, erythroblasts, 

and lymphocytes ; here and there cells suggesting the splenic cells of the 
adult organ may be seen, but giant-cells are apparently not present. A 
section taken through the splenic artery and vein with the blood contents 
of these vessels shows quite clearly that the vein contains many more 
lymphocytes than the artery, a feature which indicates that the lymphoid 
function of the spleen is already established ; whilst the presence of ery- 
throblasts in fairly large numbers in the spleen-pulp leads to the inference 
that hsemogenesis is also a splenic function at this stage — a point which 
is well established in the case of the rabbit and some other mammals. 

No Malpighian corpuscles are to be seen in a section of the organ, but in some 
places the lymphocytes seem to be more densely aggregated than in others, with an 
indication of a small artery in their neighbourhood : these masses are not, however, 
well defined. 

(vi.) The kidneys and adrenal bodies (PI. XXII. figs. 1 and 2). — At this stage the 
kidney presents histogenetic features which correspond fairly closely with those seen 
in the kidney of a four-months human foetus ; that is to say, each renal pyramid 
when sectioned in a plane passing through the cortex and the apex of the papilla is 
seen to be made up of the following parts : — ■ 

(a) A cortical zone, in the outermost layer of which are to be seen the di- 
chotomously branched endings of the tubules derived from the ingrowth 
into the metanephros of the diverticulum from the Wolffian duct ; in many 
of them the ampullary portions are continued into a coiled tubule — cut 
across many times and in various planes — the first or proximal convoluted 
tubule. At a somewhat deeper region of the cortex, the first set of 
Malpighian capsules are to be seen, these being relatively large as com- 
pared with those arising later : no sign of Henle's tube is as yet evident, 



240 MR HAROLD AXEL HAIG ON 

since no downgrowth has occurred from the convoluted tubule to form 
the loop.* 
(6) A deep zone, in which as yet connective tissue preponderates, and through 
which course the branching tubes derived from the Wolffian diverticulum ; 
these tubules possess wide lumina, and are lined by a clear cubical epi- 
thelium. They represent the rudiments of the straight and collecting 
tubules, those nearest the papilla becoming later on the ducts of Bellini ; 
in the Phocidse, according to Chievitz,! many of the secondary and tertiary 
branches of the Wolffian diverticulum disappear during development. 

The epithelium lining the convoluted tubules of the cortical zone is clear and 
cubical, the nuclei staining but feebly with basic stains : the portion of the tubule, 
however, which joins the ampulla is lined by smaller cells, the nuclei of which stain 
deeply. The glomerulus in each Malpighian capsule is a well-developed capillary 
tuft with already an indication of lobulation. 

Between the renal pyramids and supporting them there is a certain amount of 
connective tissue (columns of Bertini) in which small blood-vessels are seen cut across 
(capsular vessels of later stages). 

The adrenal bodies (PL XXII. fig. 2). — Kelatively speaking, the adrenals are much 
smaller than one would expect at this phase of development, but their histogenetic 
features are none the less instructive : each gland is enclosed in a capsule of connec- 
tive tissue in which course branches of the adrenal artery and vein, whilst smaller 
vessels (arterial) pass at right angles to the surface into the gland, being supported 
by the fine septa which are given off from the inner aspect of the capsule. 

The substance of the adrenal is made up of the following parts : — 

(a) An outer zone, the commencing zona glomerulosa, composed of folded columns 

of small cuboidal cells. 

(b) A wide intermediate zone composed of anastomosing broad columns of large 

polyhedral cells : this is the developing zona fasciculata, amongst the 
columns of which ramify the small vessels noted above as passing in from 
the capsule. The cells of this zone are characterised by their small clear 
nuclei, and their deeply-staining cytoplasm, which takes up eosin very 
readily. 
Throughout this zone are scattered small masses of rounded cells, with deeply- 
staining nuclei, the sympathetic ganglion rudiments. These are aggregated 
in larger masses in the central region of the gland, where they form the 
anlage of the medulla. In the medullary region the blood-channels are 
wide, and as yet are lined by a well-defined endothelium. 

* Chievitz (Archiv Anal. u. Embryol., Supplement, 1897) found HenLe's tubule present in an embryo of Phoca, 
145 mm. in length. The present foetus is only 122 mm. in length, 
t Op. cit. 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 241 

Compared with mammals, such as the pig, the adrenal of this Seal appears to 
be somewhat behindhand in the relative rate of its development ; thus, in a pig 
embryo of 119 mm. the medulla is well defined, and most of the sympathetic 
derivatives are confined to it alone ; but in a pig embryo of 70 mm. the histology of 
the gland is much as has been described above.* 

(vii.) The genital glands (PI. XXII. fig. 4). — As has been noted, the genital glands 
in the present instance are ovaries ; each ovary has a well-defined histological appear- 
ance, and it is possible to recognise — 

(a) An outer layer of rather high cubical epithelium. 

(b) A wide cortical zone composed of masses of rounded cells or primordial ova 

(some of these being considerably larger than others, and forming potential 
ova which will later become surrounded by a follicle of smaller cells to 
form the commencing Graafian follicles). Between the ova a good deal 
of fibro-cellular connective tissue is to be seen, and this forms a dense 
interlacing network supporting the egg-cells. 

(c) Deep to the above cortical zone comes a layer of fibro-cellular stroma, com- 

parable to the tunica albuginea of the testicle, but having a relatively 
different position in the gland. 

(d) A central portion, composed for the most part of dense stroma, with here and 

there masses of ova, which are the remains of the so-called medullary 
cords of somewhat earlier stages. 

In the mesovarium, which forms a wide peritoneal band of attachment, there are 
to be seen mesonephric tubules and glomeruli, whilst at the point of attachment of 
this band to the ovary there are some tubules, supported by stroma, forming the 
rete ovarii. 

The above histological appearances correspond closely with those to be seen in 
the ovary of a cat embryo of 94 mm. ; f no signs of developing Graafian follicles are 
to be seen, as the follicular epithelium has not as yet been formed round any of the 
larger ova. 

(viii.) The placenta (Text-fig. 2). — The details of the placenta and placentation 
have been acquired in the present instance from the study of a specimen lent to the 
author by Professor Waterston of King's College, London ; this specimen shows a 
foetus of the Weddell Seal, a closely allied species, in situ in the uterus, with the 
membranes in their proper relative position with regard to foetus and placenta. | The 
relations of the amniotic sac to the foetus and umbilical cord are such that the 
former appears to be enclosed in a small complete sac, which is quite closed towards 
the ventral aspect of the embryo, the umbilical cord upon reaching the line of 
closure dividing up into a number of branches each containing twigs from the 

* See Hertwig, op. cit. t Figured by Coert, Inaug. Dissert., Leiden, 1898. 

J Figured in Sir William Turner's Catalogue of Marine Mammals. 
TRANS. ROY. SOC. EDIN, VOL. L. PART I. (NO. 8). 34 



242 



MR HAROLD AXEL HATG ON 



arteries and vein. The amnion then spreads out on either side of the line of closure, 
the resulting folds passing anteriorly, posteriorly, and laterally to the margins of the 
placenta, over which it then spreads, covering the foetal surface of that structure. 
The branches of the umbilical arteries and vein are conducted by the above-mentioned 
folds to the margins of the placenta, and finally divide up into numerous twigs which 
enter the substance of the chorion. 

The placenta itself is of the zonary type, occupying a median zone of the uterus : 
its histology corresponds very closely with the descriptions of Grosser # and Duval 
for the zonary placentae of the cat and bitch, with the exception of certain minor 




Fig. 2. — A portion of a transverse section of the uterine wall and placenta of the 
Weddell Seal. (Semi-diagrammatic.) 



m. Muscle layers of uterine wall. 
b. Maternal blood-vessels. 
e. Epithelium lining uterine glands. 



I. Lumina of glands containing secretion (em- 
bryotrophe of Grosser). 
tr. Interglandular fastening septum. 
v. Villi of placental labyrinth. 

The foetal blood-vessels exist in considerable numbers in the villi, but are not represented in the figure ; 
the foetus contained in the above uterus had undergone about one-third of its development. 



details. Thus we find that a vertical section taken through the placenta and uterine 
wall presents the following main features (fig. 2) : — 

(a) A sheet of amnion (not shown in the figure) covering the foetal surface of the 

numerous " cotyledons" of the placenta. 

(b) The foetal portion of the placenta, composed of the rather wide laminae of the 

so-called "placental labyrinth " f between which mesodermic tissue lies; 
the sheets of this labyrinth are made up of a syncytium (derived from 
the trophoblast of earlier stages) which encloses and surrounds large 



* Vergleichende Anal, und Entvrickelungsgesch. der Eihaute u. Placenta, 1909. 
t See Grosser, op. cit. 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 243 

numbers of fetal capillaries, whilst maternal vessels of rather wider 
calibre lie in the mesodermic septa and become at times surrounded by 
portions of the syncytium. 

(c) A zone in which an invasion of the superficial gland-layer of the uterine 

mucosa has taken place, the syncytium of the villi having at an earlier 
phase converted the uterine epithelium into what Grosser terms a " sym- 
plasma," and becoming as it were welded with the mucosa at numerous 
points ; the partitions between the glands become also fixed to other 
villous tufts. 

(d) A deeper layer which comprises the bases of the uterine glands, and lies 

next the uterine muscle. The muscular coat, which is in the present case 
thin, contains large branches of the maternal blood-vessels, and these, 
where the interglandular septa pass to become fixed to the syncytium 
of the villi, pass into the mesoderm lying between the lamellae of 
the placental labyrinth. 

It appears that zone c noted above (so-called " umlagerungszone " of Strahl and 
Grosser) is, during the earlier phases, of the greatest importance in establishing the 
connection between the uterine epithelium and the syncytium of the villi ; a further 
action of the syncytium is to convert some of the decidual cells lying between the 
uterine glands into trophoblastic masses not unlike the invading syncytium itself. 

During earlier stages, stress is also to be laid upon the probability that the 
secretion of the uterine glands serves as an additional source of nutriment (" embryo- 
trophe ") to the foetus. 

From the above account it will be seen that there is a considerable histological 
similarity between the zonary placentae of the cat and Seal ; one notable difference 
is to be seen in the relatively narrow extent of the gland-layer in the uterus of the 
Seal, and another in the greater width of the laminae of the placental labyrinth. 

(ix.) The internal ear (PL XXII. fig. 6). — The semicircular canals, utricle, ampullae, 
and cochlea are well advanced in development, and lie in the cartilaginous rudiment 
of the osseous labyrinth ; the membranous labyrinth is represented by a somewhat 
thick connective tissue with a certain amount of elastic tissue entering into its 
composition. 

A transverse section across one of the semicircular canals in situ shows that the 
canal is placed very excentrically, lying against one side of the cartilaginous labyrinth, 
to which the connective tissue fixes it quite firmly. From the projecting part of the 
canal, strands or bundles of fibres pass to the opposite circumference of the cartila- 
ginous tube, and, joining here a continuation of the membrane, help to fix the canal, 
so that practically no contraction of the lumen is possible. The spaces between the 
fixing strands are filled with perilymph, whilst the canal itself is lined by a somewhat 
flattened epithelium, which later on secretes endolymph. In the utricle and ampullae 



244 MR HAROLD AXEL HAIG ON 

small elevations or " cristae" project into the lumen, and are lined by a much higher 
type of epithelium than that found in the canals ; but as yet there is no evidence of 
hair-like structures upon the free internal borders of the component cells. 

The cochlea is rather more advanced in development than would be the case in a 
three-months human foetus ; according to Krause, # the organ of Corti in the human 
foetus at birth shows that the membrana tectoria is only commencing to form, whilst 
the sensory epithelium lying upon the basilar membrane shows only a larger and a 
smaller group of columnar cells. The author's preparations of the cochlea of a three- 
months human foetus show the sensory epithelium as a group of columnar cells higher 
than the remainder in the tube, but no sign of the membrana tectoria. The scala 
tympani is present, but no membrane of Reissner as yet divides the upper cavity into 
scalse media and vestibuli, whilst the rudiments of the spiral ganglion and cochlear 
nerve are certainly to be made out, but are not at all advanced. 

In the Seal embryo under discussion, all three scalse are present, the spiral ganglion 
is a marked feature, the membrane of Reissner is well defined, and the epithelium of 
the organ of Corti is becoming differentiated, the component cells being higher upon 
the outer side, and their free borders showing a well-defined clear zone. 

(x.) The pituitary body (see text-fig. 3).- — A specimen of the pituitary gland of 
an adult Weddell Seal (Leptonychotes weddelli) was examined histologically by 
the author some time since, t and found to possess all three portions, viz. pars anterior, 
pars intermedia, and pars nervosa, highly differentiated. The Sea-Leopard Seal in its 
earlier phases of development possesses a very interesting hypophysis, which more- 
over sheds some light upon the origin of the pars intermedia. Although a complete 
account of the histogenesis of this structure is not possible in the present instance, 
there are certain points in its development which are worthy of somewhat detailed 
description, and for purposes of comparison the developing hypophyses of the rabbit 
(at the twelfth, fourteenth, and nineteenth days) and of the three-months human 
foetus were submitted to microscopical examination. 

A nearly median sagittal section taken through the pituitary of this Seal embryo 
(the gland being in situ in the sella turcica of the ossifying sphenoid bone) shows the 
following features : — 

(i.) The anterior lobe (pars anterior), consisting of irregular columns of rather 
large polyhedral cells, separated by wide blood-channels, the latter possess- 
ing a well-marked endothelial lining ; at the posterior extremity of the lobe 
the blood-channels are observed to open into large venous tributaries, which 
ultimately join up and communicate with the cavernous sinus, the latter 
being seen in section lying just anterior to the dorsum sellse of the sphenoid 
bone (fig. 3, g). 

* See Hertwig, op. cit. 

t Trans. Roy. Hoc. Kdin., vol. xlviii. part iv., No. 31, 1913. 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 



245 



(ii.) An intermediate cleft lying dorsal to the pars anterior, which is the remaining 
evidence of Rathke's pouch ; this cleft is towards its lateral aspects (not 
shown in the figure) partially filled by proliferating columns of epithelial 
cells, some of which are to be seen in the mesial section at the anterior 
extremity of the cleft. 

(iii.) The pars nervosa, or posterior lobe, lying dorsal to the cleft, and connected 
with the floor of the 3rd ventricle by the infundibulum ; part of the cavity 
of the ventricle is seen extending into the infundibulum, but this feature 
disappears during later stages to a large extent. 




Fig. 3. — A vertical section through the base of the skull to show the hypophysis cerebri in situ, lying 
in the sella turcica of the sphenoid bone. (Semi-diagrarnmatic.) 

a. Epithelium of the roof of the nasal cavity. 

b. Developing basisphenoid bone ; the ' ' irruption "-stage of ossification in cartilage is represented. 

c. Large anterior lobe of hypophysis showing cell-columns and intervening blood-vessels ; the 

dark masses are cells staining more deeply with eosin. 

d. Infundibulum, showing communication with the 3rd ventricle of brain. 

e. Posterior lobe (pars nervosa) of hypophysis. 

/. Cleft between the anterior and posterior lobes ; at the anterior end of this is seen a mass of 
cells, derived by an infolding from the anterior lobe, which will ultimately give rise to the 
pars intermedia. 

g. Cavernous sinus. 

h. Dorsum sellae of sella turcica. 



In connection with the pars intermedia, it appears that the columns of cells found 
partially filling the above-mentioned cleft are derived by a proliferation of cells at 
the upper and anterior angle of the pars anterior. Herring # describes (in the cat) 
the pars intermedia as an epithelial investment of the posterior lobe, but in the Seal, 
at least during later stages, the intermediate mass is a well-defined strip of closely 

* Joum. of Exper. Physiol., 1909. 



246 MR HAROLD AXEL HAIG ON 

packed cells, which curves round the ventral aspect of the posterior lobe and appears 
to end abruptly in a broad club-shaped extremity at the posterior and ventral margin 
of that lobe. The dorsal portion, however, passes over the upper surface of the pars 
anterior and seems to fuse with the anterior and dorsal margin of that lobe. No 
remnant of Rathke's pouch is to be seen in the fully formed pituitary of the Seal, 
whereas in the rabbit, cat, and man the pouch persists as a distinct cleft between the 
pars anterior and the pars intermedia. 

In the present instance the roof of Eathke's pouch is lined by a columnar 
epithelium, which at the anterior extremity merges into the cell-columns which later 
on fill up the pouch. Upon the dorsal aspect of the posterior lobe no epithelial 
investment is to be seen, whilst the floor of Rathke's pouch is formed by the super- 
ficial cells of the pars anterior, which are arranged in the form of an epithelium. 

The very close union maintained between the buccal and cerebral portions of the 
pituitary from the earliest stages is commented upon by Herring # and emphasised 
as having a direct bearing upon the functional importance of the gland. In the 
Seal, at the stage of development now under discussion, the union between 
the two portions is very intimate, but becomes even more marked as develop- 
ment proceeds. 

Compared with the hypophysis of a three-months human fetus, that of this Seal 
is a good deal in advance. In the human foetus at three months the connecting 
strand passing between the pars anterior and the buccal epithelium is still present, 
whilst the posterior lobe is only represented by a very narrow diverticulum from the 
floor of the 3rd ventricle. 

Again, sections taken sagittally through the pituitary of the rabbit three days 
before birth show that the size of the posterior lobe is small as compared with the 
anterior, and the cavity in the infundibulum is represented by a mere cleft lined by 
ependyma. During earlier phases in this animal (twelfth and fourteenth days) the 
relation between the size of the lobes is such that the posterior lobe is proportion- 
ately larger than during later stages. In the Seal, at the stage under discussion, the 
posterior lobe is at least one-half the size of the pars anterior ; but this relation does 
not hold in the full-grown animal, where the anterior lobe is five or six times as large 
as the pars nervosa. 

More detailed histological examination of the pituitary of the present fetus 
shows certain characters which differ from those observed in similar anatomical 
regions during later stages ; in the first place, the blood-channels in the pars anterior 
still retain their endothelial lining, a feature which disappears during development 
to a large extent, although in some parts of the full-grown anterior lobe endothelial 
cells are to be seen forming at least a partial lining to the blood-vessels. The cell- 
columns of the anterior lobe are composed of closely packed polyhedral cells, which, 
when stained with an acid stain such as eosin, may be differentiated into two 

* Op. cit. 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 247 

varieties, viz. a majority which stain lightly, and here and there isolated cells 
staining intensely ; the cytoplasm of the latter cells is also more granular in 
appearance. In the fully formed gland these deeply staining cells are more 
numerous, and collected into small groups instead of being isolated cells. The pro- 
liferating columns seen in Eathke's pouch are composed of rather cuboidal, or in 
some cases columnar, cells, which do not possess any special affinity for eosin, but 
the nuclei of which stain deeply with basic dyes ; no evidence of the syncytial 
structure seen in the pars intermedia of later stages is to be made out, nor does any 
colloid appear to have been formed as yet. The pars nervosa appears to possess 
much the same minute structure as does that part of the adult gland ; for the most 
part, neuroglia cells and fibres predominate, some of the former being spindle-shaped 
and occurring in that portion of the lobe which lies next the epithelium forming 
the roof of Eathke's pouch, their long axes being at right angles to the ventral 
surface of the lobe. The portion of the cavity of the 3rd ventricle which passes 
into the infundibulum is lined by an ependyma of the usual type, viz. rather high 
columnar epithelium, the component cells being ciliated, whilst the basal portions 
of these cells are continued as neuroglia fibres into the substance of the pars nervosa. 
There is no colloid to be detected as yet in this part of the pituitary, although in the 
fully formed gland small particles of colloidal material derived from the pars inter- 
media are to be seen scattered throughout the posterior lobe. Lying just dorsal to 
the infundibulum is a folded portion of the floor of the 3rd ventricle, which encloses 
a small fold of pia mater ; this portion later on becomes modified to form a small 
ovoid mass lying on the dorsal aspect of the posterior lobe, and contains syncytial 
strands of nucleated cytoplasm more fully described by the author in a previous 
communication.* When fully formed it is very vascular, but its functional signi- 
ficance is not at all obvious. The above somewhat brief description of some of the 
developmental aspects of the pituitary of the Sea-Leopard Seal are, of necessity, 
incomplete, on account of the fact that only a single specimen was available for 
investigation. It would be interesting to follow some of the earlier phases, inasmuch 
as the gland appears in this animal to have a high functional significance — quite as 
much so, in fact, as in some higher types. t 

General Comparative Conclusions in connection with the Anatomical and 
Histological Features presented by the Fcetus of Stenorhynchus. 

(i.) The fcetus, in the light of the above considerations, shows many points in 
common with the human foetus at the beginning of the fourth month of intra-uterine 
development. If the relative rates of growths during the earlier developmental 

* Op. cit. 

t For further details of the cytological characters of the various regions of the mammalian pituitary, see 
Schafer, " Text-Book of Microscopic Anatomy " (Quain's Anatomy, vol. ii. part i.). 



248 



MR HAROLD AXEL HAIG ON 



phases are at all comparable in the two cases, then the present foetus should have 
completed about one-third of its development. In many respects, however, this 
foetus shows an advance upon the three-months human embryo, notably in connection 
with the pituitary gland, the cerebellum, and the internal ear. 

(ii.) The other mammals (rabbit, cat, and pig) with which incidentally the foetus 
has been compared show on the whole a fairly close agreement, both anatomically 
and histologically ; one marked exception, from the anatomical point of view, is in 
connection with the kidney, which places the Seals in a small sub-group of the 
carnivora, to which the bear also belongs. 

(iii.) The developmental features of the pituitary, kidney, and brain are sufficiently 
instructive to necessitate, when possible, investigation of the earlier phases of 
development of these organs ; the acquisition of early Seal embryos is, however, a 
difficult matter, but they might with advantage be sought for, as large numbers of 
Seals are killed annually. 

In conclusion, the author wishes to thank Dr. W. S. Bruce and Professor 
Waterston for their courtesy in lending Scotia specimens of foetal Seals for investi- 
gation ; also Professor Hepburn for many valuable suggestions in connection with 
anatomical details, and Mr. T. H. Burlend, of University College, Cardiff, for much 
useful criticism concerning the arrangement of the above report. 



BIBLIOGRAPHY. 



Bailey and Miller, Text-book of Embryology, 1909. 

Bryce, T. H, Quain's Anatomy : vol. i., Embryology. 

Chievitz, Archiv Anat. u. EmbryoL, Supplement, 1897. 

Grosser, Vergleich. Anat. und Entwickelungsgeschichte der Eihaute u. Placenta, 1909. 

Haig, Trans. Roy. Soc. Edin., ibid. 

Hepburn, Trans. Roy. Soc. Edin., vol. xlviii., 1913. 

Herring, Journ. of Exper. Physiol., 1909. 

Hertwig, Handbuch der Entwickelungslehre der Wirbeltiere, 1906. 

Keibel and Mall, Text-book of Embryology, 1912. 

Schafer, E. A., Quain's Anatomy : vol. ii. part, i., Text-book of Microscopic Anatomy. 



EXPLANATION OF PLATES. 



Plate XIX. 



Fig. 1. Foetus of the Sea-Leopard Seal :- 
a.f. Anterior nipper. 
p.f. Posterior flipper. 

t. Tail. 
cl.a. Cloacal aperture. 

u. Umbilicus. 
u.c. Umbilical cord. 



am. Amnion. 
p. Placenta. 
b. Tongue : the bifid extremity showing 

between lips. 
pal.f. Palpebral fissure. 



THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 



249 



Fig. 2. Dissection of the Sea-Leopard Seal, to show the principal viscera in situ (ventral aspect): — 



1. Thyroid and lowest parathyroid. 

2. Trachea. 

3. Thymus. 

4. Left lung (upper lobe). 

5. Left auricular appendix. 
5a. Left lung (lower lobe). 

6. Ventricular portion of heart. 

7. Diaphragm. 

8. Left lobe of liver. 

9. Umbilical cord. 



10. Patent allantoic duct. 

11. Urinary bladder. 

12. Coils of small intestine. 

13. Umbilical vein. 

14. Eight lobe of liver. 

15. Gall bladder. 

15a. Right lung (middle lobe). 

16. Right auricular appendix. 

17. Right lung (upper lobe). 

18. Arch of aorta. 



Plate XX. 



Anatomical details of the brain and cranial and nasal fossae. 



Fig. 1. The brain from the dorsal aspect : — 

1. 4th ventricle (floor). 

2. Lamellae of cerebellar hemisphere. 

Fig. 2. The brain from the side : — 

1. Olfactory lobe. 

2. Fissure of Sylvius. 

3. Infundibulum. 

4. 3rd nerve. 



3. Superior surface of cerebral hemisphere. 

4. Crura cerebri. 



5. Medulla oblongata. 

6. Cerebellum. 

7. Cerebral hemisphere. 



Fig. 3. Mesial surface of a median sagittal section of head :- 



1. Lower jaw. 

2. Tongue. 

2 1 . Lamina terminalis. 

3. Hypophysis cerebri. 

4. Medulla oblongata. 

5. 4th ventricle. 

6. Cerebellum. 



7. Corpora quadrigemina. 

8. Optic thalamus. 

9. Falx cerebri. 

9 1 . Corpus callosum. 

10. Nasal fossa (lower). 

11. Nasal septum. 

12. Buccal cavity. 



Fig. 4. Nearly mesial sagittal section of brain to show distribution of grey and white matter, and general 



relations of parts : — 

1. Nuclei of vagus and hypoglossal nerves. 

2. Grey matter of cerebellar hemisphere. 

3. Pia mater. 

4. Grey matter of cortex cerebri. 

5. Superficial white matter of cortex. 

6. Grey matter lining cavity of prosen- 

cephalon, just showing. 



7. Portion of choroid plexus. 

8. Portion of 3rd ventricle in infundibulum. 

9. Grey matter in optic thalamus. 

ii. Part of prosencephalon communicating with 

lateral ventricle, 
iv. 4th ventricle. 



Fig. 5. Mesial surface of sagittal section of head, the nasal septum and falx having been removed 



t. Tongue. 

hy. Hypophysis. 

m.o. Medulla oblongata. 

cb. Cerebellum. 

p. Pons. 

TRANS. ROY. SOC. EDIN., VOL. L. PART I. (NO. 8). 



op. Optic thalamus. 

h. Cerebral hemisphere. 

olf.l. Olfactory lobe. 

eth.t. Ethmo-turbinal bone. 

mx.t. Maxillo-turbinal bone. 



35 



250 



MR HAROLD AXEL HAIG ON 



Plate XXI. 

Anatomical features of some of the viscera. 
Fig. 1. The stomach and intestines from the ventral aspect : — 



1. Coils of small intestine. 

2. Peritoneal fold attached to greater 

curvature of stomach. 

3. Pyloric end of stomach. 

4. Cardiac end of stomach. 

Fig. 2. A coronal section of the thyroid and lowest 
th. Lobules of the lateral lobe. 
p.th. Parathyroid, showing arrangement of cell 
nective tissue. 
v. Vein in capsule, with a small nerve-trunk. 

Fig. 3. The lungs from the ventral aspect : — 

a. Part of right middle lobe. 

b. Upper and lower lobes of left lung. 

c. Bronchi entering root of lung. 

Fig. 4. The retroperitoneal and pelvic viscera : — 

1. Cloacal aperture. 

2. Severed allantoic duct. 

3. Urinary bladder. 

4. Ureter (cut across). 

5. Genital duct. 

6. Left kidney. 

Fig. 5. The liver from dorsal aspect : — 
r.l. Right lobe. 
k. Surface of right lobe which lies over 
kidney. 

Fig. 6. Deep surface of the spleen : — 

a. Elongated hilus with blood-vessels. 



5. CEsophagus. 

6. First bend of duodenum, 

7. Pancreas. 

8. Part of transverse colon. 

parathyroid : — 

-columns with intervening blood-channels and con- 



d. Pulmonary vein emerging from right lung. 

e. The three lobes of right lung. 



7. Adrenal gland. 

8. Diaphragmatic ligament of mesonephros. 

9. Genital gland. 

10. Rectum, with rudimentary uterus just ventral 
to it. 



l.l. Left lobe 

l.v. Umbilical vein ( — ligamentum teres of later 
stages.) 

b. and c. Dorsal and ventral margins. 



Plate XXII. 

Figures (semi-diagrammatic) illustrating the histological features of some of the viscera. 

Fig. 1. Portion of a longitudinal coronal section of a kidney showing a quadrant of one renal pyramid 
and some intervening connective tissue : — 



a. Ampullae of terminal branches of 
Wolffian diverticulum. 
con.t. Convoluted tubule seen cut across 
several times. 
m.c. Malpighian capsules. 

Fig. 2. Part of a longitudinal coronal section of an adrenal body : — 

c. Capsule. s. Sympathetic ganglion cell masses 



w. Secondary, tertiary, etc., branches of Wolffian 
diverticulum. 
i. Intervening connective tissue between neigh- 
bouring pyramids. 



z.g. Zona glomerulosa. 
z.f. Zona fasciculata. 
m. Medulla. 



b. Blood-channels in medulla between large 
groups of sympathetic cell-masses. 



THE SYSTEMATIC ANATOMY OF A F03TAL SEA-LEOPARD. 



251 



Fig. 3. Part of a section of lung, showing the branching tubules embedded in discrete masses of con 
densed connective tissue with rudiments of muscular and elastic layers lying next the tubules. The in- 
terstitial connective tissue is also present in large relative amount. 



Fig. 4. Portion of a longitudinal section of the ovary : — 



ep. Surface epithelium (germinal epi- 
thelium). 
c. Cortical zone with primitive ova and 

some stroma. 



s. Dense stroma forming a kind of tunica 
albuginea. 
m. Medulla with a few masses of ova and 
much stroma separating them. 



Fig. 5. Part of a section of the pancreas, showing typical arrangement of branching tubular acini, with 
but little interstitial connective tissue. 

Fig. 6. Portion of a vertical section through the developing cochlea showing one turn of the spiral 
in section : — 

s.v. Scala vestibuli. 



R. Membrane of Reissner. 
ex. Canal of the cochlea. 



B. Basilar membrane, with rudimentary organ 
of corti resting upon it. 
S.T. Scala tympani. 
G. Spiral ganglion. 



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TRANSACTIONS 



OF THE 



ROYAL SOCIETY OF EDINBURGH. 

VOLUME L, PART II.— SESSION 1913-14. 




CONTENTS. 

PAGE 

IX. Stalk-eyed Crustacea Malacostraca of the Scottish 'National Antarctic Expedition. By Rev. 
Thomas R. 11. Stubbing, M.A., F.R.S., F.L.S., F.Z.S., Hon. Fellow of Worcester College, 
Oxford. Communicated by Dr J. H. Ashworth. (Plates XXIII-XXXIL), . . 253 

{Issued June 4, 1914.) 

X. The Aborigines of Tasmania. Part III. The Hair of the Head compared with that of other 
Ulotrichi and, with Australians and Polynesians. P.y Principal Sir William Turner, 
K.C.B., D.C.L., F.E.S., Knight of the Royal Prussian Order Pour le Merite, Emeritus 
Professor of Anatomy. (With Figures in Text.), ..... 309 

(Issued June 30, 1914.) 

XL The Pinna-Trace in the Ferns. By R. C. Davie, M. A., B.Sc, late Robert Donaldson Research 
Scholar in the University of Glasgow, Lecturer in Botany in the University of Edinburgh. 
Communicated hy Professor I. Bayley Balfour, F.R.S. (Plates XXXIII-XXXV.), ~ . 349 

(Issued August 18, 1914.) 

XII. Studies on the Pharmacological Action of Tetra- Alkyl- Ammonium Compounds. II. The 

Action of Tetra- Ethyl- Ammonium Chloride. By Professor C. R. Marshall. (With 

Figures in Text.), ........ .379 

(Issued July 29, 1914.) 

XIII. Rods from Gough Island, South Atlantic (collected by the Scottish National Antarctic Expedi- 

tion, 1902-1904). By Robert Campbell, M.A., D.Sc, Lecturer in Petrology in the 
University of Edinburgh. Communicated by Professor James Geikie, D.C.L., LL.D., F.R.S. 
(Plate XXXVI.), .......... 397 

(Issued October 3, 1914.) 

XIV. On a New Species of Sclerocheilus, with a Revision of the Genus. By J. II. Ashworth, D.Sc, 

Lecturer in Invertebrate Zoology in the University of Edinburgh. (Plate XXXVII, and 

Four Text-figures.), . . . . . . . . .405 

(Issued Jidy 2, 1915.) 



XV. Atlantic Sponges collected by the Scottish National Antarctic Expedition. By Jane Stephens, 
B.Sc. Communicated by Dr W. S. Bruce. (Plates XXXVIII-XL.), 

(Issued May 5, 1915.) 



423 



XVI. On the Fossil Osmundacex. By R. Kidston, LL.D., F.R.S., F.G.S., Foreign Mem. Imper. 
Mineralogical Society of Petrograd, Hon. Sec. R.S.E. ; and D. T. Gwynne-Vaughan, M.A., 
F.R.S.E., M.R.I. A., Professor of Botany, University College, Reading. (Plates XLI-XLIV.), 469 

(Issued December 9, 1914.) 

XVII. Studies on the Pharmacological Action of Tetra- Alkyl- Ammonium Compounds. III. The 

Action of Methyl-Ethyl-Ammonium Chlorides. By Professor C R. Marshall, . .481 

(Issued February 26, 1915.) 



EDINBURGH: 

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( 253 ) 



IX. — Stalk-eyed Crustacea Malacostraca of the Scottish National Antarctic 
Expedition. By the Rev. Thomas R. R. Stebbing, M.A., F.R.S., F.L.S., 
F.Z.S., Hon. Fellow of Worcester College, Oxford. Communicated by Dr 

J. H. ASHWORTH. 

(MS. received December 2, 1913. Read March 16, 1914. Issued separately June 4, 1914.) 

[Plates XXIII.-XXXIL] 

As the Scotia was engaged in marine research throughout the whole of its long 
voyage out and home, it was only to be expected that Antarctic specimens would 
form but a small fraction of the result. In like manner, as several of the halting- 
places, such as the Cape of Good Hope and the Falkland Islands, had been visited 
by many keen naturalists on earlier occasions, it was quite likely that species of the 
group with which this report is concerned would have been in numerous instances 
already observed. Accordingly, out of some fifty species here discriminated, only 
six claim to be new, and not more than ten can be regarded as Antarctic or sub- 
Antarctic in their place of capture. In the vast extension, however, which marine 
zoology has for some time past been receiving, some retardation in the stream of 
discovery may not be unwelcome to the systematist. Familiar forms which would 
otherwise amply repay a thorough reinvestigation are apt to be thrust on one side, 
when striking novelties are for ever appealing to be introduced. This expedition, 
like others before it, affords fair evidence that the Macrura are both abundant and 
varied in great depths of the ocean. But only too often the frailty of their fabric 
leaves them in a tantalisingly mutilated condition when they reach the surface. 

CONSPECTUS OF THE SPECIES. 

BRACHYURA. 
Brachyura genuina. 
Tribe OXYRRHYNCHA. 

Family Inachid^e. 

Achzeopsis thomsoni (Norman), Gough Island, 40° 20' S., 9° 56' W. 
Coryrhynchus algicola, n. sp., 18° 24' S., 37° 58' W. 

Eurypodius latreillii, Guerin, Falkland Islands, and Burdwood Bank, 54° 25' S., 
57° 32' W. 

TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 9). 36 



254 RKV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

Family Mamaiid.e. 
Macrocceloma concavus, Miers, 18° 24' S., 37° 58' W. 

Family Parthenopid^e. 
Lambrus verrucosus, Studer, off Pyramid Point, Ascension Island. 

Tribe CYCLOMETOPA. 

Family Xanthid^e. 
Xanthodius parvulus (Fabricius), Porto Grande, St Vincent. 

Family Portunice. 
Lupa sayi, Gibbes, from Gulf weed, Sargasso Sea. 

Family Corystid^e. 
Nautilocorystes ocellatus (Gray), entrance to Saldanha Bay, South Africa. 

Tribe CATOMETOPA. 

Family GoneplaciD;E. 
Goneplax angulatus (Pennant), off Dassen Island, near Saldanha Bay, South 

Africa. 
Pilumnoplax heterochir (Studer), Gough Island, 40° 20' S., 9° 56' W. 

Family Grapsid^e. 

Grapsus maculatus (Catesby), St Helena, St Vincent, and Ascension Island. 
Cyclograpsus punctatus, Milne-Edwards, False Bay, South Africa. 
Planes minutus (Linn.), from Gulf weed, Sargasso Sea. 
Plagusia capensis, de Haan, Saldanha Bay and Cape Town. 
Percnon planissimus (Herbst), Porto Grande, St Vincent. 

Family Gecarcinid^e. 
Gecarcinus lagostoma, Milne-Edwards, Ascension Island. 

Family Hymenosomatid.e. 

Hymenosoma orbicularis, Desmarest, Saldanha Bay, South Africa. 
Halicarcinus planatus (Fabricius), Falkland Islands and South Orkneys. 

Tribe OXYSTOMATA. 

Family Calappid^e, 
Mursia cristimanus, de Haan, off Dassen Island, Saldanha Bay, South Africa. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 255 

Brachyura anomala. 

Family Dromiid^. 

Dromia dormia (Linn.), 48° 06' S., 10° 05' W. 

Dromia sp. ?, Saldanha Bay, South Africa. 

Pseudodromia latens, Stimpson, Saldanha Bay, South Africa. 

Family Latreilliid^e. 
Latreillia elegans, Roux, off Pyramid Point, Ascension Island. 

Macrura anomala. 

Tribe PAGURIDEA. 

Family Lithodid.e. 

Lithodes antarcticus, Jacquinot, Burdwood Bank, 54° 25' S., 57° 32' W. 
Paralomis granulosus (Jacquinot), Burdwood Bank, 54° 25' S., 57° 32' W. 

Family Pagurid^e. 

Pagurus arrosor (Herbst), Cape Town. 

Pagurus calidus, Risso, Ascension Island. 

Eupagurus forceps, Milne-Edwards, Falkland Islands. 

Ewpagurus modicellus, n. sp., Pyramid Point, Ascension Island. 

Calcinus tcdismani, A. Milne-Edwards and Bouvier, Porto Grande, St Vincent. 

Tribe GALATHEIDEA. 

Family Galatheid^e. 

Munida subrugosus (White), Falkland Islands. 
Munida gregarius (Fabricius), juv., Falkland Islands. 

Tribe HIPPIDEA. 

Family Albuneid^e. 
Albunea guerinii, Lucas, St Helena. 

Macrura genuina. 
Tribe SCYLLARIDEA. 

Family Scyllarid^e. 
Scyllarides elisabethse (Ortmann), off St Helena. 



256 REV. T. R. R. STEBB1NG ON STALK-EYED CRUSTACEA MALACOSTRACA 

Family Palinurid^e. 
Jasus lalandii (Milne-Edwards), Saldanha Bay, South Africa. 

Tribe PEN.ELDEA. 

Family Pen^eid^e. 

Gennadas kempi, n. sp., 39° 48' S., 2° 33' E. 
Gennadas parvus (?), Bate, 48° 00' S., 9° 50' W. 

Family Leuciferid^;. 
Petalidium foliaceus, Bate, 48° 00' S., 9° 50' W. 

Tribe CARIDEA. 

Family Paljsmonid^e. 

Leander squilla (Linn.), Porto Grande, St Vincent. 

Leander affinis (Milne-Edwards), Saldanha Bay, South Africa. 

Leander tenuicornis (Say), from Gulf weed, Sargasso Sea. 

Family Hippolytid^. 

Hippolyte acuminatus, Dana, from Gulf weed, Sargasso Sea. 
Latreutes fucorum (Fabricius), from Gulf weed, Sargasso Sea. 
Nauticaris brucei, n. sp., Gough Island, 40° 20' S., 9° 50' W. 
Nauticaris magellanicus (A. Milne-Edwards), Falkland Islands. 

Family Pasiph^id^e. 

Phye scotix, n. sp., 68° 32' S., 12° 49' W. ; 71° 22' S., 16° 34' W. 
Phye rathbunw, n. sp., 48° 00' S., 9° 50' W. 

Family Nematocarcinid^e. 

Nematocarcinus lanceopes, Bate, 71° 22' S., 16° 34' W. ; and (?) 39° 48' S., 
2° 33' E. 

SCHIZOPODA. 

Order MYSIDACEA. 
Sub-order LOPHOGASTRIDA. 

Family Eucopiid,e. 
Eucopia sp., 39° 48' S., 2° 33' E. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 257 

STOMATOPODA. 

Family Squillid^e. 

Squilla armatus, Milne-Edwards, off Dassen Island, South Africa. 
Lysioerichthus edwardsii (Eydoux and Souleyet), juv., 19° 59' N., 22° 34' W. 
Though not properly coming within the scope of this report, the occurrence of 
the following species may for convenience' sake be mentioned here : — 
Bopyrina latreuticola (Gissler), from Gulf weed, Sargasso Sea. 
Lanceola sestivus, Stebbing, 68° 32' S., 10° 52' W., surface. 
Nebalia bipes (Fabricius), Saldanha Bay, South Africa, 25 fathoms. 

BRACHYURA 
Brachyura genuina. 

Tribe OXYRRHYNCHA. 

The classification here adopted is substantially the same as that used in the 
General Catalogue of South African Crustacea, to be found in the Annals of the 
South African Museum, vol. vi. part iv., issued in 1910. 

Family Inachid.e. 

Genus Achaeopsis, Stimpson. 

1857. Achaeopsis, Stimpson, Pr. Ac: Sci. Philad., vol. ix. p. 219. 

1873. Dorynchus, Norman, in Wyville-Thomson's Depths of the Sea, p. 174, fig. 34. 

1880. Lispognathus, A. Milne-Edwards, Etudes Crust, reg. Mexicaine, p. 319. 

1886. Achseopsis, Miers, Rep. Voy. " Challenger," vol. xvii. part xlix. p. 18. 

1886. Lispognathus, Miers, Rep. Voy. " Challenger," vol. xvii. part xlix. p. 27. 

1886. ,, Perrier, Explorations sous-marines, p. 298. 

1893. Achseopsis, Ortmann, Zool. Jahrb., vol. vii. p. 36. 

1910. ,, Stebbing, Annals S. Afr. Mus., vol. vi. part iv. p. 285. 

1910. Dorynchus, Stebbing, Annals S. Afr. Mus., vol. vi. part iv. p. 285. 

1911. Achseopsis, Rathbun, Tr. Linn. Soc. London, vol. xiv. part ii. p. 247. 

Miers states that Achseopsis is distinguished from Inachus " merely by having 
the postocular as well as the prseocular spine distinctly developed, and by the more 
or less falciform dactyli of the three posterior ambulatory legs." From A. spinulosus, 
Stimpson, 1857, he says that his own A. giintheri, 1879, is distinguished " by having 
but a single very long perpendicular spine on the gastric region." Like these, A. 
superciliaris , Ortmann, 1893, has the rostrum not deeply divided. From Inachus 
and Achaeopsis Miers considered Lispognathus " distinguished by the well-developed 
rostral spines." For that genus Perrier notices as also characteristic the long 
slender ambulatory legs. Miss Rathbun, however, in 1911, without discussion, sinks 
Lispognathus, and by inference also Dorynchus, as synonyms of the earlier Achseopsis. 



258 REV. T. R. R. STEBBfNG ON STALK-EYED CRUSTACEA MALACOSTRACA 

The generic name Dorynchus depended not on a definition but on the dorsal view of 
a specimen. The word Lispognathus, signifying " a smooth jaw," is not very appro- 
priate for Norman's species, in which the large third joint of the external maxillipeds 
is well studded with tubercles. Further, it has been shown by Doflein that in respect 
of length and divergence the rostral spines of Dorynchus thomsoni are very variable, 
so that its removal to Achseopsis is made the less difficult, and two rival claimants 
to the generic name may conveniently be dispensed with. 

Achseopsis thomsoni (Norman). 

1873. Dorynchus thomsoni, Norman, Depths of the Sea, p. 174, fig. 34. 

1880. Lispognathus furcillatus, A. Milne-Edwards, Crust. Mexic, p. 349, pi. xxxiA. fig. 4. 

1886. Lispognathus thomsoni, Miers, Rep. Voy. " Challenger;" vol. xvii. part xlix. p. 28, pi. v. fig. 2. 

1886. ,, ,, Perrier, Explor. sous-marines, p. 298, fig. 218. 

1900. „ ,, A. Milne-Edwards and Bouvier, Crust. Decap. " Travailleur-Talisman," 

p. 148, pi. iii. fig. 8, pi. xxi. figs. 8-14. 
1904. ,, ,, Doflein, Ergebn. Deutschen Tiefsee-Exp., vol. vi. p. 75. 

1908. ,, ,. Hansen, Ingolf-Exp., "Crust. Malac.," vol. iii. part ii. p. 11. 

1910. Dorynchus thomsoni, Stebbing, Annals S. Afr. Mus., vol. vi. part iv. p. 286. 

1911. Achseopsis thomsoni, Rathbun, Tr. Linn. Soc. London, vol. xiv. part ii. p. 247. 

A. Milne-Edwards and Bouvier in 1900 still regard L. furcillatus as specifically 
distinct (loc. cit., p. 148), and on pp. 151, 152 (twice by a slip calling it L. furcatus) 
say that " L. thomsoni is distinguished from L. furcillatus by the more slender form 
of the carapace, narrow front, rostral spines less divergent, spines of carapace more 
marked." Doflein with ample material decides that the differences are based on a 
young specimen. Miss Rathbun says of an adult male example taken at Seya de 
Malha from a depth between 300 and 500 fathoms: "This example has parallel 
horns about a quarter as long as remainder of carapace. It differs from typical 
specimens in having the anterior gastric and anterior branchial spines obsolete or 
reduced to low tubercles. The species is very close to A. spinulosus, Stimpson 
(Smithson Misc. Coll., xlix., 1907, p. 21, pi. iii. figs. 5, 5a), which has shorter legs, 
described as minutely spinulous above, but there is no indication, in description or 
figure, of the terminal spine on the merus joints. A. spinulosus is an inhabitant 
of shallower water (10 fathoms in Simon's Bay, Cape of Good Hope)." As sug- 
gested above, the conspicuously long legs of A. thomsoni seem to separate it de- 
cisively from the other species of the genus. But if A. spinulosus has anything 
like the variability established for A. thomsoni, the validity of A. gilntheri must be 
regarded as resting on a very insecure foundation. 

The Scotia specimens have a denticle at the middle of each of the divergent 
rostral horns and a larger subdistal one visible from below. The eyes have a 
tubercle on the peduncle and another subdistal on the corneal portion. In the 
first antennae the third joint of the peduncle is oval, wider than the preceding joint; 
the principal fiagellum carries numerous filaments. The palp of the mandibles 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 259 

appears to be membranous, the first joint scarcely distinct from the second, the 
third joint carrying two or three setae. In the third maxillipeds the inner margin 
of the third joint is finely but irregularly denticulate, the surface tubercles not closely 
set. The eggs of a small female are numerous, not very small. 

Locality.— Gough Island, lat. 40° 20' S., long. 9° 56' 30" W. ; depths 75 and 100 
fathoms ; April 21-23 and 24, 1904. 

The extraordinary distribution of this species has been noted by several authors, 
extending as it does from the Faroes to the West Indies, the Cape of Good Hope, 
Australia, and the Indian Ocean, with depths varying from 106 m. to 1326 m. Fuller 
details are given by Milne-Edwards and Botjvier, and by Doflein. 

Gen. Coryrhynchus, Kingsley. 

1860. Podonema (preocc), Stimpson, Ann. Lyceum N.H. New York, vol. vii. p. 19. 
1870. „ Stimpson, Bull. Mus. Comp. Zool., vol. ii. p. 126. 

1879. Podochela (part), A. Milne-Edwards, Miss. Sci. Mexique, "Crust.," part v. p. 189. 

1879. Podonema, Miers, J. Linn. Soc. London, vol. xiv. No. 79, p. 643. 

1879. Coryrhynchus, Kingsley, Pr. Ac. Sci. Philad., p. 384. 
1886. ,, (subgen.), Miers, Rep. Voy. " Challenger" vol. xvii. part xlix. p. 11. 

1900. ,, (subgen.), Young, West Indian Stalk-eyed Crust., p. 13. 

1901. Podochela (part), Rathbun, Bull. U.S. Fish. Comm.for 1900, vol. ii. p. 53. 

The peculiarity of an almost circular rostrum in species which otherwise clearly 
belong to the Oxyrrhyncha or sharply rostrate crabs may be taken to justify the 
separation of the genus from its near ally Podochela. 

Coryrhynchus algicola, n. sp. 
Plate XXIII. 
This new species is approximate to C. riisei (Stimpson), 1860, and C. spatulifrons 
(A. Milne-Edwards), 1879. It agrees with the latter in the short broad form of the 
rostrum, the former being distinguished from both by having the rostrum longer 
and narrower. On the other hand, the more pronounced angles of the distally 
widened fourth joint in the third maxillipeds here agree with C. riisei, and not with 
C. spatulifrons. In both of those species the fingers of the chelipeds are described 
by the French author as finely denticulate, and he figures those of his own species 
to correspond with that statement. The new species has the edges of these fingers 
not dentate but crenulate, the alternate projections of one finger neatly fitting the 
hollows of the other. Here also a pair of tubercles occur on the surface of the 
carapace between the eyes, which would seem to be absent from the other species 
under comparison. It may be judged from the copious supply of hooked and other 
setae with which the whole exposed surface is furnished in all three species, that 
all adopt similar methods of concealment. The Scotia specimen was a mere garden 
of seaweed, and the limbs were far more ready to leave the body than either body 
or limbs were to part with the investing weeds. The pellucid rostrum looked like 
a bit of weed, and one of the long second pereeopods was so thickly matted that it 



260 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

had ceased to be in any way suggestive of a limb. The eyes have an apical tubercle 
carrying a setule. The first and second joints in the palp of the mandible seem 
to be in coalescence. The pleon of the female forms a deep almost circular bowl, 
the last three segments in coalescence constituting more than half of it. Length of 
carapace 19 mm., greatest breadth 13 mm., second perseopod about 47 mm. long. 
For C. spatulifrons the corresponding measurements are 19, 14, and 44 mm. 
Locality.— Lat. 18° 24' S., long. 37° 58' W. ; depth 36 fathoms ; Station 81. 

Genus Eurypodius, Guerin. 

1828. Eurypodius, Guerin, Mem. Mus. Hist. Nat. Paris, vol. xvi. pp. 349, 350. 

1900. „ Stebbing, Pr. Zool. Soc. London, p. 527. 

1910. „ Rathbun, Pr. U.S. Mus., vol. xxxviii. pp. 571, 612. 

Eurypodius latreillii, Guerin. 

1828. Eurypodius latreillii, Guerin, Mem. Mus. Hist. Nat. Paris, vol. xvi. p. 354, pi. xiv. 

1900. „ „ Stebbing, Pr. Zool. Soc. London, p. 527. 

1905. Eurypodius lalreillei, Lagerberg, Schwed. Sudpol. Exp., vol. v. part vii. p. 17 (with copious 

synonymy). 
1910. Eurypodius latreillii, Rathbun, Pr. U.S. Mus., vol. xxxviii. pp. 571, 612. 

The study of this rather abundant species by numerous authors has led to its 
receiving a considerable variety of names. These are latreillia (probably an error of 
the press), tuberculatus, audouinii, andouinii (error of the press), cuvieri, septen- 
trionalis, brevipes, latreillei (probably meant for a correction), danse. All are now 
regarded as synonyms of the original E. latreillii, Guerin. In Lagerberg's useful 
list of the synonymy the Crustacea of the Coquille are attributed to H. Milne- 
Edwards, by mistake for Guerin, who naturally uses the form latreillii, not the 
erroneous latreillia, which only reappears in Gay's Hist. Chile, 1849, where Nicolet 
is pleased to refer it to Guerin' s own writings. According to the last-named author, 
the colour in fresh (condition is greenish brown. The discussions instituted by 
various authors make it fairly certain that the supposed specific differences depend 
on sex and age or inconstant variation. Lagerberg had at his disposal specimens 
varying in the length of the carapace between 8 and 87 mm., with a breadth of 
4 '5 mm. for the smallest and of 65 mm. for the largest. The Scotia specimens do 
not reach beyond a medium size. 

Localities. — Falkland Islands, Port Stanley, from 2 and 4 fathoms ; Port William, 
from 6 fathoms ; Station 346, Burdwood Bank, lat. 54° 25' S., long. 57° 32' W., depth 
56 fathoms. According to Lagerberg, the greatest depth at which this species has 
hitherto been obtained is the 70 fathoms recorded by Miers in his Challenger report. 
Eurypodius longirostris, Miers, was dredged in 175 fathoms. 

Family Mamaiid/E. 

1905. Mamaiidx, Stebbing, Gilchrist's Mar. Invest. S.A. Crust., part iii. p. 22 ; and Pr. Biol. Soc. 
Washington, vol. xviii. p. 157. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 261 

Genus Macrocceloma, Miers. 

1879. Macrocceloma, Miers, J. Linn. Soc. London, vol. xiv. p. 665. 

1886. ,, Miers, Rep. Voy. " Challenger," vol. xvii. part xlix. p. 79. 

1901. „ Rathbun, Bull. U.S. Fish. Oomm.for 1900, vol. ii. p. 73. 

Macrocceloma concavus, Miers. 

1886. Macrocosloma concava, Miers, Rep. Voy. " Challenger" vol. xvii. part xlix. p. 81, pi. x. 

figs. 2, 2a-b. 
1898. Macrocceloma concavum, Rathbun, Pr. U.S. Nat. Mus., vol. xxi. p. 576. 
1901. „ „ Rathbun, Bull. U.S. Fish. Comm. for 1900, vol. ii. p. 75. 

In this species the carapace is deeply concave upon the hepatic regions, " the 
spines of the rostrum are short, in the adult less than one-fourth the length of the 
carapace, they are nearly straight, divergent, and separate by a triangulate inter- 
space." The arrangement of the tubercles on the somewhat damaged carapace of 
the Scotia specimen appears also to answer the description given by Miers, and the 
third maxillipeds and limbs agree with his account and illustrations, with this excep- 
tion, that in our specimen the finely denticulate inner margins of the fingers in the 
chelipeds leave a gap only quite close to the hinge. The length of the carapace 
measured from the base of the rostral spines is 29 mm., the spines themselves 
5 mm., the breadth at the eyelobes 20 mm., and at the widest part to the rear 
23 mm., not including the lateral spines which are here outstanding. 

Locality.— -Lat. 18° 24' S., long. 37° 58' W. ; depth 36 fathoms; Station 81, 
December 20, 1902. 

For the gender of the specific name see Knoivledge, vol. xxxiii., No. 504, p. 259 ; 
and No. 509, p. 470, 1910. 

Family Parthenopice. 

1834. " Parthenopiens " (tribe), Milne-Edwards, Hist. Nat. Crust., vol. i. p 347. 

1847. Parthenopidee, White, List Crust. Brit. Mus., p. 41. 

1901. „ Rathbun, Bull. U.S. Fish. Comm. for 1900, vol. ii. p. 79. 

Genus Lambrus, Leach. 

1815. Lambrus, Leach, Tr. Linn. Soc. London, vol. xi. pp. 308, 310. 

1895. „ Alcock, J. Asiat. Soc. Bengal, vol. lxiv. p. 259. 

1901. „ Rathbun, Bull. U.S. Fish. Comm. for 1900, vol. ii. p. 79. 

Lambrus verrucosus, Studer. 

1882. Lambrus verrucosus, Studer, Abhandl. K. Ak. Wiss. Berlin, vol. ii. p. 9, pi. i. figs. 2a, 2b. 
1886. „ „ Miers, Rep. Voy. " Challenger," vol. xvii. part xlix. p. 93. 

The species is notable for the very deep furrows which separate the tuberculate 
median ridge of the carapace from its tuberculate branchial regions. The spinulose 
rostrum is more produced than in Sttjder's figure, and the chelipeds are more 
verrucose and dentate in the Scotia specimen than in that which Studer represents. 
The eyes show minute warts both on and below the cornea. The species belongs to 

TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 9). 37 



262 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

that section of the genus which Miers defines as having the fourth joint of the 
ambulatory limbs more or less spinose or tuberculated. All the six segments of the 
pleon are also warty. 

Locality. — Off Pyramid Point, Ascension, from depth of 40 fathoms ; June 10, 
1904; Station 507. 

Tribe CYCLOMETOPA. 
Family XanthidtE. 

1898. Xanthidx, Alcock, Jour. Asiat. Soc. Bengal, vol. lxvii. part ii. p. 69. 
1910. ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 296. 

The family is here taken in the wider sense. 

Genus Xanthodius, Stimpson. 

1859. Xanthodius, Stimpson, Ann. Lye. Nat. Hist. New York, vol. vii. p. 52. 
1901. „ Rathbun, Bull. U.S. Fish Comm. for 1900, vol. ii. p. 27. 

A. Milne-Edwards, who gives the reference to Stimpson as on p. 6, thinks that 
Stimpson distinguished Xanthodius from Leptodius on indifferent grounds, the crest 
on the endostome in the former being incomplete, and often occurring in a recognised 
species of the latter genus. Later authorities, however, appear to agree in upholding 
Stimpson's determination, though Miss Rathbun does not insist on the character to 
which A. Milne-Edwards refers, and Dr Young speaks of (subgenus) Leptodius 
as having " endostome without trace of longitudinal carinse. 

Xanthodius parvulus (Fabricius). 

1793. Cancer parvulus, Fabricius, Ent. Syst., vol. ii. p. 451. 

1858. Chlorodius americanus, Saussure, Mem. Soc. Hist. Nat. Geneve, vol. xiv. p. 430 (14), pi. i. fig. 5. 

1860. Xanthodius americanus, Stimpson, Ann. Lye. Nat. Hist. New York, vol. vii. p. 209 (81). 

1879. Leptodius americanus, A. Milne-Edwards, Miss. Sci. Mexique, part v. p. 269. 

1897. Xanthodius parvulus, Rathbun, Ann. Inst. Jamaica, vol. i. No. i. p. 15. 

1900. Leptodius (Xanthodius) americanus, Young, West Indian Stalk-eyed Crust., p. 147. 

1901. Xanthodius parvulus, Rathbun, Bull. U.S. Fish Comm. for 1900, vol. ii. p. 27. 

1908. „ „ Verrill, Tr. Connect. Ac. Sci., vol. xiii. p. 340, text-fig. 12, pi. xiv. fig. 4. 

Miss Rathbun's identification of de Saussure's species with the Cancer 
parvulus of Fabricius was based on " types examined." The description given by 
Fabricius is therefore inexact, since he states the front to be entire, whereas it is 
notched in the centre ; he speaks of the sides as tridentate, but they are quadriden- 
tate, and he says that the feet are short, smooth, with fingers black at the apex. As 
he is no doubt referring to the chelipeds, only the colour note is accurate, while the 
limbs are not specially short, and have the wrist and hand rough, " eroded in reticulat- 
ing lines." The thumb of the larger cheliped in the Scotia male specimen has a 
large blunt tooth on the inner margin near the hinge, and two smaller teeth near the 
apex ; the movable finger has a curved and hollowed apex and a median tooth on 
the inner margin. The fourth joint of the third maxillipcds is short and broad. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 263 

The ambulatory limbs are short, with setose fingers. The pleon is five-segmented. 
The measurements of the specimen coincide with those given by de Sattssure, length 
14 mm., breadth 22 mm., width of front 6 mm. 

Locality. — St. Vincent, Porto Grande, shore N.E. ; December 1, 1902 ; Station 24. 

Family Portunid^e. 

Genus Lupa, Leach. 

1813. Lupa, Leach, Edinb. Encycl., pol. vii. p. 390. 

1833. Neptunus, de Haan, Crust. Japonica, decas 1, p. 7. 

1897 Portunus, Rathbun, Pr, Biol. Soc. Washington, vol. xi. pp. 155, 160. 

1908. Lupa, Stebbing, Annals S. Afr. Mus., vol. vi. part i. p. 11. 

Several other references for the genus may be obtained from those given for the 
following species. From the neighbouring and very similar genus Callinectes, 
Stimpson, the present is distinguished in the male sex by having the pleon triangular, 
without the narrow ending which in Callinectes gives it as it were the shape of the 
capital letter T. 

Lupa sayi, Gibbes. 

1802. Portunus pelasgicus, Bosc, Hist. Nat. Crust., vol. i. p. 219, pi. v. fig. 3 (Portune pelasgique). 

1817. Lupa pelagica, Say, J. Ac. Sci. Philad., vol. i. p. 97. 

1830. Portunus pelasgicus, Bosc and Desmarest, Hist. Nat. Crust,, vol. i. p. 235, pi. v. fig. 3 (Portune 

pelagique). 
1850. Lupa sayi, Gibbes, Pr. Amer. Assoc, p. 178. 

1852. ,, „ Dana, U.S. Expl. Exp., vol. xiii. part i. p. 273, pi. xvi. fig. 8. 
1861. Neptunus sayi, A. Milne-Edwards, Arch. Mus. Hist. Nat., vol. x. p. 317, pi. xxix. fig. 2. 
1878. „ „ A. Milne-Edwards, Crust. Mexic, p. 210. 

1886. Neptunus (Neptunus) sayi, Miers, Rep. Voy. " Challenger," vol. xvii. part xlix. p. 173. 

1897. Portunus sayi, Rathbun, Ann. Lnst. Jamaica, vol. i. p. 22. 

1898. „ „ Rathbun, Pr. U.S Mus., vol. xxi. p. 592 

1908. „ „ Verrill, TV. Connect. Ac. Sci., vol. xiii. pp. 373, 374, 376, text-fig. 25, pi. xviii. 

fig. 2, pi. xxi. fig. 1. 

It is probable that this species was included by Herbst in his Cancer pelagicus, 
Krabben und Krebse, vol. i. p. 159, 1783, and represented by his fig. 55 on pi. viii. 
Miers remarks that " the convex, marbled carapace, short lateral epibranchial spines, 
and the absence of a spine on the posterior margin of the merus [fourth joint] of the 
chelipedes are characteristic of this species." He observes that " a large series of 
specimens of this common pelagic species was taken from the Gulf weed" by the 
Challenge!^ Expedition. The larger of two female specimens obtained by the Scotia 
measured 66 mm. from point to point of the epibranchial spines, with a length of 
32 mm. for the carapace. A male specimen similarly measured 70 x 38 mm. The 
pleon in this agreed with Dana's single figure, which represents this part of the 
organism. The male organs are very slender except at the base, and their apices, 
which curve strongly outwards, nearly reach the end of the telson. 

Locality.— From Gulf weed, 29° 54' N., 34° 10' W. to 33° 53' N., 32° 27' W. ; 
$ June 30, 1904, ? June 29 and July 1, 1904 ; Stations 537, 538, and 539. 



264 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

Family Corystid^e. 

1899. Corystidee, Alcock> Jour. Asiat. Soc. Bengal, vol. lxviii. pp. 5, 103. 

Genus Nautilocorystes, Milne-Edwards. 

1833. Dicera (preoccupied), de Haan, Crust. Japon., decns 1, pp. 4, 14. 
1837. Nautilocorystes, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 149. 
1900. „ Stebbing, Gilchrist's Mar. Invest. S. Afr., vol. i. p. 16 (with synonymy). 

Nautilocorystes ocellatus (Gray). 

1831. Corystes ocellata, Gray, Zool. Miscellany, vol. i. p. 39. 

1833. Corystes {Dicera) 8-dentata, de Haan, Crust. Japon., decas 1, p. 15. 

1837. Nautilocorystes ocellatus, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 149; and later in the 

undated plates of the Regne animal, pi. xxiii. figs. 2, 2a-c. 
1843. Dicera 8-dentata, Krauss, Siidafrik. Crust., p. 27. 
1847. Nautilocorystes ocellatus, White, List Crust, in Brit. Mus., p. 53. 
1857. ,, ,, Stimpson, Pr. Ac. Sci. Philad., p. 39. 

1900. Nautilocorystes octodentatus, Stebbing, Gilchrist's Mar. Invest. S. Afr., vol. i. p. 17. 
1907. Nautilocorystes ocellatus, Stimpson, Smithson. Misc. Coll., vol. xlix. p. 89 [N. octodentatus 

(de Haan) in editor's footnote]. 
1910. Nautilocorystes octodentatus, Stebbing, Annals S. Afr. Mus., vol. vi. part iv. p. 311. 

My attention has been recently called by Dr Calman to the first of these 
references, which I had overlooked. The oversight was perhaps excusable, seeing 
that Milne-Edwards prints the name Nautilocorystes ocellatus as if the species 
were his own, and White, who gives reference to Gray, gives it without date after 
his reference to Milne-Edwards ! 

Locality. — Entrance to Saldanha Bay, South Africa ; depth 25 fathoms ; 
Station 483. 

Tribe CATOMETOPA. 

Family Goneplacid^e. 

1900. Gonoplacidse, Alcock, J. Asiat. Soc. Bengal, vol. lxix. p. 282. 
1910. Goneplacidse, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 312. 

Genus Goneplax, Leach. 

1814. Goneplax, Leach, Edinb. Encycl., vol. vii. p. 430. 
1817. „ Latreille, Regne animal, vol. iii. p. 16. 

Goneplax angulatus (Pennant). 

1777. Cancer angulatus, Pennant, British Zoology, vol. iv. p. 7, pi. v. fig. 10. 
1910. Goneplax angulata, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 312. 

The Scotia specimens were taken May 18, 1904. 

Locality. — Off Dassen Island, near Saldanha Bay ; depth 35 fathoms ; Station 480. 

Genus Pilumnoplax, Stimpson. 

1858. Pilumnoplax, Stimpson, Pr. Ac. Philad., vol. x. p. 93. 

1910. ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 315. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 265 

Pilumnoplax heterochir (Studer). 

1882. Pilumnus heterochir, Studer, Abhandl. K. Ak. Wiss. Berlin, part ii. pp. 6, 14, pi. i. fig. 3, a-d. 
1886. Pilumnoplax heterochir, Miers, Rep. Voy. " Challenger," vol. xvii. part xlix. p. 227, pi. xix. fig. 1, a-d. 

Miers and Studer both call attention to the peculiarity in this species that the 
palm of the larger cheliped is smooth except at the base, while in the smaller 
cheliped the palm externally is covered with numerous granules. The fingers of 
both limbs retain their dark colour in spirit. Studer says that in life the colour is 
intense orange-red, with only the fingers of the chelae black. 

Locality. — Gough Island, depths 75 and 100 fathoms; the lower depth recorded 
on April 22, 1904, the other between April 21 and 23 ; Stations 460 and 461. 

Family Grapsid^e. 
1900. Grapsidas, Alcock, J. Asiat. Soc. Bengal, vol. lxix. part ii. pp. 283, 389. 

Genus Grapsus, Lamarck. 

1801. Grapsus, Lamarck, Syst. Anim. sans Vertebres, p. 150. 
1900. ,, de Man, Mem. Soc. Zool. France, vol. xiii. p. 48. 
1910. „ Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 317. 

Grapsus maculatus (Catesby). 

1758. Cancer grapsus, Linn., Syst. Nat., ed. 10, p. 630. 

1771, 1743 (reissue 1771). Pagurus maculatus, Catesby, Nat. Hist. Carolinas, vol. ii. p. 36, pi. xxxvi. 

fig. 1. 
1908. Grapsus grapsus, Verrill, Tr. Connect. Ac. Sci., vol. xiii. p. 317, pi. x. fig. 6; pi. xi. fig. 2 (with 

synonymy). 
1910. Grapsus maculatus, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 317. 

According to Alcock, the carapace of a large specimen is 64 mm. long and 68 mm. 
broad. In the largest of the Scotia specimens the carapace is 32'5 mm. long and 
38 mm. broad, with a front 14 mm. broad. 

Localities. — St Helena, James Bay, June 1 and 2, 1904; Station 499. Ascension 
Island, June 7, 1904 ; Station 507. Two small specimens from the shore, Porto 
Grande, St Vincent, December 1, 1902, Station 24, appear to belong to this species. 

Genus Cyclograpsus, Milne-Edwards. 
1837. Cyclograpsus, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 77. 

Cyclograpsus punctatus, Milne-Edwards. 

1837. Cyclograpsus punctatus, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 78. 

1838. Gnathochasmus barbatus, M'Leay, Annulosa of S. Africa, p. 65, pi. iii. 
1910. Cyclograpsus punctatus, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 318. 

Locality. — Specimens of this prettily marked species, in full agreement with 
M'Leay's coloured figure of it, were obtained by the Scotia Expedition from the shore 
of False Bay, May 8, 1904 ; Station 479. 



266 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

Genus Planes, Bowdich. 

1825. Planes, Bowdich, Excursions in Madeira and Porto Santo, p. 15, figs. 2a, 2b. 
1910. ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 320. 

This genus is placed by Alcock in his division Varuninse. 

Planes minutus (Linn.). 

Plate XXIV. 

1758. Cancer minutus, Linn., Syst. Nat., ed. 10, vol. i. p. 625. 
1825. Planes clypeatus, Ex. in Madeira, etc., p. 15, figs. 2a, 2b. 
1905. Planes minutus, Stebbing, Gilchrist's Mar. Invest. S.A. Crust., part iii. p. 43. 
1908. ,, ,, Verrill, Tr. Connect. Ac. Sci., vol. xiii. p. 325, text-fig. 7, pi. xiii. fig. a-f, 

pi. xxvii. fig. 6. 

A very large number of specimens were obtained by the Scotia from the Gulf 
weed on June 29 and 30 and July 1, 1904. In most cases the colouring is lost, but 
in some instances there is a well-marked pattern of dark brown on a light ground. 
In this species the third maxillipeds have the fourth joint much shorter than the 
third, but quite as broad, the outer margin strongly rounded, the fifth joint inserted 
in an excavation of the front margin of the fourth, and, when folded down, not 
reaching beyond that joint's inner edge. The principal joint of the exopod does not 
reach the end of the endopod's fourth joint. The specimen figured was rather per- 
plexingiy distinguished by retaining as preserved a dark-brown colour and showing 
oblique striae on both .sides of the carapace, while also the indent below the antero- 
lateral angles was exceptionally well marked. 

Localities. — Saldanha Bay, South Africa, 25 fathoms ; Station 483. And also from 
29° 54' N., 34° 10' W., to 33° 53' N., 32° 27' W. ; Stations 537, 538, and 539. 

Genus Plagusia, Latreille. 
1806. Plagusia, Latreille, Gen. Crust, et Insect., vol. i. p. 33. 
1900. ,, Alcock, J. Asiat. Soc. Bengal, vol. lxix. pp. 297, 436. 

1905. ,, Stebbing, Marine Invest. S. Afr. Crust., part iii. p. 46. 

1908. ,, Verrill, Tr. Connect. Ac. Sci., vol. xiii. p. 332. 

1910. ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 322. 

Immature Form. 

1852. Marestia, Dana, U.S. Expl. Exp., vol. xiii. p. 487. 

1910. ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv., p. 348. 

From the essential agreement of the mouth-organs in the adult and Megalopa 
specimens it seems clear that the form which Dana assigned to his genus Marestia 
is really a juvenile stage of Plagusia. Both forms occur abundantly together. In 
his full account of the genus Alcock invites especial attention to a feature of the 
third maxillipeds, that " their exognath has no flagellum." But this must be taken 
with some limitation, since de Ha an figures a small flagellum for his Plagusia 
(Icnhjics, and I find one in P. capensis. It is slender, without setse except at the 
apex, the apex in the Megalopa stage indicating faintly a second joint. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 267 

Plagusia capensis, de Haan. 

Plate XXVIc. 

1835. Plagusia capensis, de Haan, Crust. Japon., decas 2, pp. 31, 58. 

1837. Plagusia tomentosa, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 92. 

1905. Plagusia capensis, Stebbing, Marine Invest. S. Afr. Crust., part iii. p. 47. 

1910. Plagusia chabrus, Rathbun, Pr. U.S. Mus., vol. xxxviii. pp. 591, 616. 
1910. ,, ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 322. 

The reasons which I gave in 1905 for upholding the name given by de Haan to 
this species still seem to me valid, while its identification with the obscurely described 
Cancer chabrus of Linn^us appears to be quite arbitrary. 

Locality. — Saldanha Bay, Station 483 ; and at the coaling jetty, Cape Town 

docks, Station 478. 

Genus Percnon, Gistel. 

1835. Acanthopus (preoccupied), de Haan, Crust. Japon., decas 2, p. 29. 

1848. Percnon, Gistel, Naturg. Thierreichs, p. viii. 

1900. „ Rathbun, Pr. U.S. Mus., vol. xxii. p. 281. 

1910. „ Stebbing, Ann. S. Afr. Mus., vol. vi. pt. iv. p. 324. 

Percnon planissimus (Herbst). 

1804. Cancer planissimus, Herbst, Krabben u. Krebse, vol. iii. part iv. p. 3, pi. lix. fig. 3. 

1825. Plagusia clavirnana, Desmarest, Consid. gen. Crust., p. 127, pi. xiv. fig. 2. 

1838 (?) „ „ Milne-Edwards, Regne anim., "Undated Crust.," pi. xxiii. fig. 3. 

1900. Liolophus planissimus, Alcock, J. Asiat. Soc. Bengal, vol. lxix. p. 439 (copious synonymy). 

1902. ,, ,, de Man, Abhandl. Senckenberg. not. Gesellsch., vol. xxv. part iii. p. 543, 

pi. xx. fig. 12. 
1900. Percnon planissimum, Rathbun, Pr. U.S. Mus., vol. xxii. p. 281. 

1908. ,, ,, Verrill, Tr. Connect. Ac. Sci., vol. xiii. p. 334, pi. x. fig. 3, pi. xii. fig. 4. 

1910. „ „ Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 324. 

This strikingly marked little species has an extensive distribution, being recorded 
from the East Indies, Australia, New Zealand, the Cape of Good Hope, Chile, Japan, 
and the Bermudas. According to Krauss the South African specimens have the 
carapace dusky greenish brown, with a bright blue stripe down the middle, and the 
feet striped (gebandert) with reddish brown and yellow. Alcock for the Indian 
species says that " the colour in life is dark green, the nude streaks being bright 
green." Herbst's figure shows the rostrum projecting strongly beyond the flanking 
lobes, which in the Scotia specimen as in the figures by Desmarest, Milne-Edwards, 
and Verrill nearly or quite equal the rostrum in length. Here also, as in Verrill's 
figure, several of the legs are cross-banded instead of being longitudinally striped. 

Locality. — St Vincent, Porto Grande, shore ; December 1, 1902 ; Station 24. 

Family Gecarcinid^e. 

1837. " Gecarciniens," Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 19 
1852. Gecarcinidse, Dana, U.S. Erpl. Exp., vol. xiii. p. 374. 
1894. ,, Ortmann, Zool. Jahrb., vol. vii. p. 732. 



268 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

Genus Gecarcinus, Leach. 
1814. Gecarcinus, Leach, Edinb. Encycl., vol. xvii. p. 430 (see also pp. 391, 435). 
1825. ,, Desmarest, Consid. gen. Crust., p. 113. 

1835. Ocypoda, Freminville, Ann. Sci. Nat., ser. 2, vol. iii. p. 224. 
1836 (?) Gecarcinus, Guerin, Icon. Regne Anim., "Crust." pi. v. 
1837. „ Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 25. 

1858. ,, Saussure, Crust. Mexi que- Antilles, p. 23. 

1886. Geocarcinus, Miers, Rep. Voy. " Challenger," vol. xvii. part xlix. p. 217. 

1893. Gecarcinus, Stebbing, History of Crustacea, Internat. Sci. Ser., vol. lxxiv. p. 79. 

1894. „ Ortinann, Zool. Jahrb., vol. vii. pp. 732, 740. 

1895. ,, Bouvier, Bull. Nat. Hist. Mus. Paris, vol. i. p. 8. 
1897. ,, Rathbun, Ann. Inst. Jamaica, vol. i. No. 1, p. 24. 
1901. „ Rathbun, Bull. U.S. Fish. Comm. for 1900, vol. ii. p. 14. 
1910. „ Rathbun, Pr. U.S. Mus., vol. xxxviii. p. 812. 

In this genus of land crabs the small terminal joints of the endopod of the third 
maxillipeds, sometimes called the palp, are completely concealed in a ventral view by 
the large fourth joint, and the angle which this joint forms with the third leaves 
open a lozenge-shaped gap between the two members of this pair of maxillipeds. 

In 1837 Milne-Edwards distinguished three species based respectively on the 
Cancel' ruricola of Linnaeus and many other authors, the Ocypoda lateralis of 
Freminville, and an Australian form which he named G. lagostoma. His new 
species agreed with G. ruricola in having the fingers of the ambulatory limbs armed 
with six rows of spine-teeth instead of the four which characterise G. lateralis. Its 
carapace was said to be smaller than that of G. ruricola, which, however, is not the 
case. The colour is not mentioned, so that the only point of distinction was left 
dependent on the third maxillipeds, in which the merus or fourth joint is described 
as having a deep fissure on its internal edge above the following joint, while in 
G. ruricola there is no notable fissure. 

Gecarcinus lagostoma, Milne-Edwards. 

1837. Gecarcinus lagostoma, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 27. 

1886. Geocarcinus lagostoma, Miers, Rep. Voy. "Challenger," vol. xvii. part xlix. p. 218, pi. xviii. 

figs. 2, 2a-c. 
1893. „ „ Benedict, Pr. U.S. Mus., vol xvi. p. 537. 

1893. Gecarcinus lagostoma, Ortmann, Decap. und Schizop. Plankton-Exp., p. 58. 

1894. „ ,, Ortmann, Zool. Jahrb., vol. vii. p. 740. 
1900. „ „ Rathbun, Pr. U.S. Mus., vol. xxii. p. 277. 

Two large female specimens of this species from Ascension Island were given to 
the Scotia collection by Mr Chalmers. They fully equal in size the specimen from 
the Challenger Expedition figured by Miers, and surpass it in the massiveness of the 
chelipeds. Mr Chalmers also presented a much smaller female, matching in size 
and colouring a male specimen taken by the naturalists of the Scotia on June 1, 
1904. These small specimens, after nearly nine years in preservative liquid, still 
have the larger part of the carapace dark purple with six light spots, one outside 
each eyelobe, the other two pairs at spaces converging towards the median groove. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 269 

The colouring might easily suggest such names as the violet or purple crab, which 
have been applied to G. ruricola, and shows correspondence with the figure which 
Guerin gives of G. lateralis. Miers says of the small specimen from Bermuda 
which he identifies with G. lagostoma, that in colour it nearly resembles specimens 
of G. lateralis, a species to which Heilprin and Ortmann definitely assign it, only 
with the difference that Ortmann regards G. lateralis as merely a young form of G. 
ruricola. The latter opinion is not accepted by Miss Rathbun, and does not appear 
to be tenable, if reliance can be placed on the number of spine-rows in the ambulatory 
fingers as having specific value. These rows are six in number in the small as well 
as in the large specimens of G. lagostoma. But in a specimen from Antigua, sent 
me by G. R. Forrest, Esq., which measures 58 mm. in breadth by a length of 45 mm., 
these rows are limited to four. The specimen has a very large left cheliped, with 
gaping fingers, and by the pleon and appendages is clearly an adult male. A third 
pair of carinse on the ambulatory fingers is at least indicated, though it is not spini- 
ferous. Also the fourth joint of the third maxillipeds has a fissure, though less in 
extent than that in G. lagostoma. On the whole, it is perhaps not surprising that 
authorities have varied in applying the names ruricola, lateralis, and lagostoma 
with others which are noted by Miers, Ortmann, and Rathbun. 

The male specimen obtained by the Scotia has the pleon appendages wide apart, 
about reaching the end of the sixth segment, with a short subapical fringe directed 
outward. Breadth of carapace 43 mm., length 34 mm. 

Locality. — Ascension Island, 5 to 18 fathoms; Station 507. 

Family Hymenosomatid^e. 

1858. Hymenosomidx, Stimpson, Pr. Ac. Sci. Philad., vol. x. p. 108. 

1905. Hymenosomatidx, Stebbing, Gilchrist's Mar. Invest., "S.A. Crust.," part iii. p. 49. 

1906. Hymenosomidx, W. H. Baker, TV. R.S. South Australia, vol. xxx. p. 112. 
1910. Hymenosomatidae, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 331. 

As genera belonging to this family there have been named Hymenosoma, 
Desmarest, 1825; Elamena, Milne-Edwards, 1837; Halicarcinus, White, 1846; 
Hymenicus, Dana, 1852. The specific names which have been distributed among 
these genera need to be revised by someone who can compare specimens from the 
several localities concerned. 

Genus Hymenosoma, Desmarest. 

1825. Hymenosoma, Desmarest, Consid. gen. Crust., pp. xvii, 163. 

1900. „ Stebbing, Pr. Zool. Soc. London, pp. 520, 523. 

1905. ,, Stebbing, Gilchrist's Mar. Invest., "S.A. Crust.," part iii. p. 49. 

1907. ,, Stimpson, Smithson. Misc. Coll., vol. xlix. p. 144. 

1910. ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 331. 

In 1900 I was inclined to agree with Professor Haswell's opinion that the 
separation of Halicarcinus from Hymenosoma rested " on extremely slight points of 
TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 9). 38 



270 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

distinction." But at least in the typical species of each genus the third maxillipeds 
are very unlike, the third and fourth joints being about twice as long as broad in 
Hymenosoma orbicularis, whereas the length and breadth are subequal in Hali- 
carcinus planatus ; besides that the details of shape are not the same. A further 
dissimilarity concerns the ringers of the walking-legs, which in H. orbicularis have 
the inner margin of the fingers smooth, but clothed almost to the tip with two dense 
rows of long plumose setae inserted on either side. In H. planatus these ringers 
have the inner margin fringed with spine-teeth alternating with setse of very 
moderate length. In the figures and description which Mr W. H. Baker supplies in 

1906 for Hymenosoma rostratus, Haswell, he represents a maxilliped agreeing with 
that of H. planatus, and says that the dactyli of the ambulatory legs " are slightly 
curved and carry a series of small teeth of about equal size with hairs between." Thus 
the species seems unsuitable for union with Hymenosoma. Professor Chilton in 

1907 decides that Hymenosoma depressus, Jacquinot, belongs to the genus in which 
Jacquinot placed it, and though Jacqtjinot's figure of the external maxilliped is 
obviously untrustworthy, the limbs in Chilton's specimen had the other mark of the 
genus in " the fact that the terminal joints of the last four pairs of legs were fringed 
with hairs and looked as if they were used as swimming organs." 

Hymenosoma orbicularis, Desmarest. 

Plate XXVa. 

1825. Hymenosoma orbiculare, Desmarest, Consid. gen. Crust., p. 163, pi. xxvi. figs. 1, la-e. 

1837. „ „ Milne-Edwards, Hist. Nat. Crust, vol. ii. p. 36. 

1838. (?) „ ,, Milne-Edwards, Regne anim., illustr., pi. xxxv. figs. 1, la. 
1858. Hymenosoma geometricum, Stimpson, Pr. Ac. Philad., p. 108 (54). 

1904. Hymenosoma orbiculare, Doflein, Deutsch. Tiefsee-Exp., vol. vi. p. 88. 

1905. Hymenosoma geometricum, Stebbing, Gilchrist's Mar. Invest., "S.A. Crust.," part iii. p. 50. 
1907. Hymenosoma geometricum and Hymenosoma orb ictdare, Stimpson, Smithson. Misc. Coll., vol. xlix. 

p. 144. 
1910. Hymenosoma orbiculare and Hymenosoma geometricum, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. 

pp. 331, 332. 
1913. Hymenosoma orbiculare, var. geometricum, Balss, Schultze, Forschungsreise in Siid Afrika, vol. v. 

part ii. p. 113 (64). 

It may seem presumptuous to cancel Stimpson's species, but it was founded on a 
single small specimen, and his remark that the " feet are sparsely provided with fine, 
inconspicuous hairs," may have been due to a quite accidental condition. Desmarest 
gives no indication of the fringe of short setae with which the broad end of the 
female pleon is beset. His figure of the external maxillipeds shows the third joint 
much smaller than the fourth, but this misleading error is corrected by Milne- 
Edwards in the illustrated edition of the Regne animal. In the chelipeds of this 
species the hand is much more swollen in the male than in the female, and the 
movable finger of the male has a tooth on the inner margin which appears to vary 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 271 

considerably in its expansion and denticulation. Among the specimens' obtained by 
the Scotia the largest, a male taken on the shore, Houtjes Bay, May 19, 1904, has 
a carapace 20 mm. long by 15 mm. broad ; the next largest, a female, has the cara- 
pace 19 mm. in length, including the rostrum as in the other instance, with a breadth 
of 15 mm. at the widest part of the granulated rim ; but, in both examples, if the 
bulging lateral parts are included, the apparent breadth becomes at least equal to the 
length, in good accord with the specific name orbicularis. The short narrow pleon of 
the male is in striking contrast to the greatly expanded pleon of the female. 

Localities. — Entrance to Saldanha Bay, depth 25 fathoms, May 21, 1914 ; Station 
483. Within Saldanha Bay on May 19, on the shore, and in depths of 5 and 8 to 10 
fathoms, and the largest female specimen on May 20 ; Station 482. 

Genus Halicarcinus, White. 

1846. Halicarcinus, White, Ann. Nat. Hist., ser. 1, vol. xviii. p. 178. 

1900. „ Stebbing, Pr. Zool. Soc. London, p. 521. 

1907. ,, Stimpson, Smithson. Misc. Coll., vol. xlix. p. 145. 

Further references to the genus will be deducible from the synonymy of the 
following species : — 

Halicarcinus planatus (Fabricius). 

1775. Cancer planatus, Fabricius, Syst. Ent., p. 403. 

1846. Halicarcinus planatus, White, Ann. Nat. Hist., ser. 1, vol. xviii. p. 178., pi. ii. fig. 1. 

1891. „ „ Mocquard, A. Milne-Edwards, Miss. Cap Horn, "Crust.," p. 27. 

1893. ,, ,, Ortmann, Zool. Jahrb., vol. vii. p. 31. 

1900. ,, ,, Stebbing, Pr. Zool. Soc. London, p. 524, pi. xxxvi.B. 

1902. „ „ Hodgson, " Southern Cross " Exp., p. 231. 

1903. ,, ,, Ortmann, Princeton Exp. Patagonia, vol. iii. p. 660. 

1904. „ „ Doflein, Deutsch. Tiefsee-Exp., vol. vi. p. 87. 

1905. ,, ,, Lagerberg, Schwed. Sudpol. Exp., vol. v. part vii. p. 25. 

1909. ,, ,, Chilton, Subantarctic Is. N. Zealand, p. 609. 

1910. ,, „ Eathbun, Pr. U.S. Mus., vol. xxxviii. p. 570. 

Numerous specimens were obtained by the Scotia at the Falkland Islands, in 
January 1903, in Port Stanley Harbour and in shore pools at Cape Pembroke, and 
again on January 31, 1904, on the shore at Port William. In this last gathering, as in 
the earlier one from shore pools, the male specimens were distinguished by the con- 
siderable size of the chelipeds. A single specimen is labelled November 1903, 
Macdougal Bay, South Orkneys. 

Tribe OXYSTOMATA. 

1841. Oxystomata, de Haan, Crust. Japonica, decas 5, p. 111. 

Family Calappid^e. 
1851. Calappidx, Dana, U.S. Expl. Exp., vol. xiii. pp. 390, 393. 



272 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

Genus Mursia, Desmarest. 

1825. Mursia, Desmarest, Consid. gen. Crust., p. 108. 

1900. ,, Stebbing, Gilchrist's Mar. Invest., "S.A. Crust.," part i. p. 21 (with synonymy). 

1904. „ Doflein, Deutsch. Tiefsee-Exp., vol. vi. p. 36. 

1906. „ Rathbun, Bull. U.S. Fish. Comm. for 190S, part iii. pp. 887-8. 

1911. ,, Rathbun, Tr. Linn. Soc. London, vol. xiv. part ii. p. 198. 

Doflein discusses the various species that had been assigned to the genus down 

to 1904. 

Mursia cristimanus, de Haan. 

1837. Mursia cristimanus, de Haan, Crust. Japonica, decas 3, p. 70. 

1839. Mursia cristimana, de Haan, Crust. Japonica, decas 4, p. 73, pi. E (mouth-organs). 

1900. Mursia cristimanus, Stebbing, S.A. Crust., part i. p. 22 (with synonymy). 

1904. Mursia cristimana, Doflein, Deutsch. Tiefsee-Exp., vol. vi. p. 38, pi. xvi. fig. 5-12, pi. xviii. fig. 1. 

To distinguish this species from M. armatus, de Haan, and its varieties or near 
allies, Doflein relies especially on three characters : the front of the carapace produced 
to a sharp central tooth ; the hind margin smooth, without teeth, only slightly 
undulating and carrying a fine row of granules ; and thirdly, the under margin of the 
chelae beset with coarse teeth, not exceeding eight or nine in number. With these 
characters the Scotia specimen agrees. 

For determining the correct specific name the synonymy given by Dr Doflein 
introduces some perplexity. He cites M. cristimana, Latreille, from Cuvier, 
Regne animal, ed. 2, p. 39, 1829, but the name cristimana does not occur in that 
work, so far as I can find, either on p. 39 of vol. iv. or elsewhere. On the other 
hand, Doflein supplies a reference which I omitted in 1900, namely, Mursia cristata, 
Milne-Edwards, in Atlas zu Cuvier, Regne animal, ed. 3, Taf. xiii. fig. 1 and la, to 
which he attaches the date 1836. If that date can be depended on, the name of the 
species should be cristatus instead of cristimanus, since the latter is not earlier 
than 1837, so far as known. It is true that in 1837 Milne-Edwards gives a reference 
to pi. xiii. of the 3rd edition of the Regne animal, but it does not follow that the 
undated plate had been published a year earlier, so that de Haan's cristimanus may 
still hold its ground, till proof is forthcoming of the true date of publication for 
pi. xiii. in the 3rd edition of the Regne animal (see additional note, p. 307). 

Locality. — Off Dassen Island, South Africa, from depth of 35 fathoms, May 18, 
1904 ; Station 480. 

Brachyura anomala. 

1893. Brachyura anomala (part), Stebbing, Hist. Crust., Internat. Sci. Ser., vol. Ixxiv. p. 133. 
1899. ,, „ Alcock, Deep-Sea Brachyura Investigator, p. 6. 

1910. „ „ Stebbing, Ann. S.A. Mus., vol. vi. p. 341. 

Family DromiidtE. 

1899. Dromiidx, Alcock, Journ. A.S. Bengal, vol. lxviii. pp. 128, 135. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 273 

Genus Dromia, J. C. Fabricius. 
1798. Dromia, J. C. Fabricius, Suppl. Ent. Syst., p. 359. 

Dromia dormia (Linn.). 

1763. Cancer dormia, Linn., Amoen. Acad., vol. vi. p. 413. 

1910. Dromia dormia, Stebbing, Ann. S. Afr. Mus., vol. vi. p. 342. 

A large specimen of what I suppose to be this species was obtained by the Scotia 
in lat. 48° 06' S. s long. 10° 05' W., the depth reached by the trawl being 1742 fathoms, 
at Station 451, May 13, 1904. 

Some small specimens, taken on the shore, Houtjes Bay, Saldanha Bay, belong to 
this family, but their position in it remains for the present indeterminate. 

Genus Pseudodromia, Stimpson. 
1858. Pseudodromia, Stimpson, Pr. Ac. Sci. Philad., vol. x. p. 226 (64). 

Pseudodromia latens, Stimpson. 

1858. Pseudodromia latens, Stimpson, Pr. Ac. Sci. Philad., vol. x. pp. 226, 240. 

1910. „ „ Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 340 (with references). 

Locality. — Saldanha Bay, South Africa, from 5 fathoms, May 20, 1904 ; Station 482. 

Family Latreilliid^e. 

1899. Latreillidce, Alcock, Journ. Asiat. Soc. Bengal, vol. lxviii. part ii. pp. 130, 165. 
1902. Latreilliidm, Stebbing, Gilchrist's Mar. Invest., "S.A. Crust.," part ii. p. 23. 

Genus Latreillia, Roux. 

1828. Latreillia, Roux, Crust. Mediterranee, part v. pi. xxii. 

1894. Latreillea, A. Milne-Edwards and Bouvier, Res. Camp. Sci. Monaco, fasc. vii. p. 59. 

Latreillia elegans, Roux. 

1828. Latreillia elegans, Roux, Crust. Mediterranee, pi. xxii. 

1902. ,, „ Stebbing, S.A. Crust., part ii. p. 24 (with synonyms). 

The small specimens obtained by the Scotia indicate that the relations of length 
between the fingers and palm of the chelipeds are variable. The feathering of the 
penultimate joint of the fifth perseopod is well shown. It is highly desirable that 
specimens of this remarkable organism should be preserved separately, if possible, 
when first captured, as otherwise they shed their limbs with vexatious freedom, 
the sharp prickles on the long, stiffly geniculating joints tending to hopeless entangle- 
ment with extraneous objects. 

Locality. — Off Pyramid Point, Ascension, from a depth of 45 fathoms, June 10, 
1904 ; Station 507. 



274 REV. T. R. K. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

MACRURA. 
Macrura anomala. 

1893. Macrura anomala, Stebbing, Hist. Crust., Internat. Sci. Ser., vol. lxxiv. p. 147. 
1901. „ ,, Alcock, Catal. Indian Deep-Sea Crustacea, p. 204. 

1908. Heieromacrura, Verrill, Tr. Connect. Ac Sci., vol. xiii. p. 433. 

1910. Macrura anomala, Stebbing, Ann. S. Afr. Mus., vol. vi. part. iv. p. 349. (From this reference 
other references may be traced.) 

Tribe PAGURIDEA. 

1901. Paguridea, Alcock, Catal. Indian Deep-Sea Crust., p. 205. 
1905. ,, Alcock, Catal. Indian Decap. Crust., part ii. p. 1. 

Family Lithodid^;. 

1853. Lithodidx, Dana, U.S. Expl. Exp., vol. xiii. p. 1430. 

1900. „ Stebbing, Pr. Zool. Soc. London, p. 530. 

1901. ,, Alcock, Catal. Indian Deep-Sea Crust., p. 231. 

Genus Lithodes, Latreille. 

1806. Lithodes, Latreille, Gen. Crust, et Insect., vol. i. p. 39. 

1905. „ Stebbing, Gilchrist's Mar. Invest., "S.A. Crust.," part iii. p. 69. 

1913. „ Balss, Abhandl. K. Bayer. Ah. Wiss., suppl., vol. ii. part ix. p. 73. 

Lithodes antarcticus, Jacquinot. 

1843-1847. Lithodes antarctica, Jacquinot, Voy. Pole Sud, Atlas, pi. vii., pi. viii. figs. 9-14. 

1847. Lithodes antarcticus, White, Crust. Brit. Mus., p. 56. 

1849. Lithodes antarctica, Nicolet, Gay's Hist. Chile, "Zool.," vol. iii. p. 182. 

1852. ,, „ Dana, U.S. Expl. Exp., vol. xiii. p. 427, pi. xxvi. fig. 15. 

1853. ,, „ Lucas, Voy. Pole Sud, "Zool.," vol. iii. p. 90. 

1891. ,, ,, A. Milne-Edwards, Mission Cap Horn, "Crust.," p. 24. 

1905. Lithodes antarcticus, Lagerberg, Schwed. Sudpol. Exp., vol. v. part vii. p. 12 (giving many other 

important references). 
1910. Lithodes antarctica, Rathbun, Pr. U.S. Mus., vol. xxxviii. p. 595. 

Nicolet says that the vernacular name of this fine crustacean is Centolla, that 
it is much appreciated as food, not only by human beings, but also by seals, which 
carry the creatures ashore and dash them against the rocks to get at the meat 
without being inconvenienced by the strong spines of the carapace. The dried 
carapace, he says, is hung up by the peasants in their cottages to act as a 
barometer, by its reddening for fine weather and becoming pale for approaching 
rain. He gives the length as 7f inches, with a stretch of the limbs reaching 29^ inches. 
Dana says : " Specimens are often 5 inches long, with a breadth of 4^ inches, 
the longest legs being 9^ inches long. The exuvia of one, procured by us, was 
8 inches in length, with the longest legs 15 inches in length." According to 
Miklouho-Maclay (quoted by Lagerberg), at the Isle of Chiloe this crustacean 
is known as Barometro Araucano, because " the ordinary colour of the shell 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 275 

during dry weather is light grey, which as soon as the air gets damp becomes 
gradually covered with spots of a dark (reddish) tint. The increase of humidity 
in the atmosphere makes the spots larger, so that the shell is at last quite of a 
dark (reddish) colour." If this and Nicolet's contrary account are both true, one 
can only infer that weather forecasts in those regions are as little to be trusted as 
in our own climate. 

The specimen obtained by the Scotia had the characteristic rostrum distally 
upturned, with two pairs of lateral teeth ; between these there is a small central 
tooth ; the peduncles of the eyes are denticulate, and a large tooth on the outer 
side of each orbit projects a little beyond the eye ; the pleon is practically 
symmetrical, without appendages, but with a medio-ventral process as shown in 
Jacquinot's fig. 12. 

Length of carapace, including the rostrum, 49 mm., greatest breadth 44 mm. ; 
length of fourth perseopods 110 mm. ; of the third not shorter. 

Locality.— Station 346. Burdwood Bank, lat. 54° 25' S., long. 57° 32' W. ; depth 
56 fathoms; December 1, 1903. 

Together with the above was taken a small pellucid specimen, unfortunately with 
rostrum broken off, measuring 18 mm. in breadth of carapace, by a length of 22 mm., 
allowing for the lost rostrum. As with the larger specimen, the pleon of this juvenile 
indicated that it was of the male sex. 

Genus Paralomis, White. 

1856. Paralomis, White, Pr. Zool. Soc. London, vol. xxiv. p. 134. 

1900. „ Stebbing, Pr. Zool. Soc. London, p. 531. 

1908. „ Hansen, Lngolf-Ezp., " Crust. Malac," vol. iii. part ii. pp. 22, 24. 

1913. „ Balss, Abhandl. K. Bayer. Ah. Wiss., suppl., vol. ii. part ix. p. 76. 

The validity of this generic name has been questioned by Benedict, who in 1894 
regarded it as a synonym of Echinocerus, White, 1848. 

In 1847, in the List of Crustacea in the British Museum, White named without 
defining a genus Echidnocerus, with a species E. cibarius. In the following year, in 
the Proc. Zool. Soc, London, p. 47, pis. ii., iii., he described and figured the form in 
question as a new species and subgenus, giving the first name as Echinocerus in the 
text, but as Echidnocerus on the plates. It may perhaps be assumed that Echinocerus 
was a misprint. However that may be, Benedict adopted it for Jacquinot's 
Magellanic species, and instituted a new genus Leptolithodes for Henderson's Para- 
lomis aculeatus, 1888, and several others. Subsequent opinion, however, has made 
Leptolithodes, and not Echidnocerus, a synonym of Paralomis. But that some of 
these generic divisions are not very easy to follow may be inferred from the remarks 
which Hansen makes in 1908 when describing a new species, Paralomis bouvieri. 
After stating that " the marginal plates on the third abdominal segment are quite 
free in the male, but quite fused with the lateral plates in the female," he adds, 
" as this feature in the marginal plates of the third segment is generally considered 



276 REV. T. E. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

an important generic character, the male should be referred to Acantholithus, 
Stimps., the female to Paralomis. I have preferred to place the species with the 
latter, as it shows some resemblance to P. aculeatus, Hend." 

Paralomis granulosus (Jacquinot). 

1843-1847. Lithodes granulosa, Jacquinot, Voy. Pole Sua 1 , Atlas, pi. viii. figs. 15-21. 
1856. Paralomis granulosa, White, Pr. Zool. Soc. London, vol. xxiv. p. 134. 
1900. „ „ Stebbing, Pr. Zool. Soc. London, p. 532. 

1903. „ ,, Ortmann, Princeton Exp. Patagonia, vol. iii. p. 658. 

1905. ,, ,, Lagerberg, Schwed. Siidpol. Exp., vol. v. part vii. p. 14. 

The Scotia specimens of this tolerably abundant species are small, the carapace 
of the larger, a female, being 36 mm., of the smaller only 23 mm., in length and 
breadth. The variations which develop in specimens of different sizes are discussed 
by Lagerberg, who had at his disposal six specimens, the smallest measuring 17 
mm. by 17 '5 mm., while the largest, a male, had a carapace 90 mm. long and 95 mm. 
broad. He figures the last four joints of the right male chela as measuring 
133 mm. in length and displaying very conspicuous teeth on the wrist. In the 
Scotia female specimen these teeth are very prominent but less irregular. Lager- 
berg records 100 m. as the greatest hitherto known depth for the occurrence of this 
species, which is practically in agreement with the Scotia record. 

Locality. — Burdwood Bank, lat. 54° 25' S., long. 57° 32' W. ; depth 56 fathoms 
or 102 m. ; December 1, 1903 ; Station 346. 

Family Pagurid^e. 
1852. Paguridse, Dana, U.S. Expl. Exp., vol. xiii. p. 435. 

Pagurus arrosor (Hcrbst). 
1908. Pagurus arrosor, Stebbing, Ann. S. Afr. Mus., vol. vi. part i. p. 22. 
Under the above reference I have discussed the synonymy and interesting 
ornamentation of this species. 

Locality. — A characteristic specimen, in a shell of the genus Triton, was obtained 
by the Scotia from coaling jetty, No. 1, Cape Town docks, May 14, 1904 ; Station 477. 

Pagurus calidus, Bisso. 

1826. Pagurus calidus, Risso, His. Nat. Eur. Merid., vol. v. p. 29. 

1888. ,, „ Henderson, Rep. Voy. " Challenger," vol. xxvii. part lxix. p. 57. 

1905. ,, ,, Alcock, Indian Decap. Crust., part ii. fasc. i. p. 170 (ample synonymy). 

This handsomely coloured species is strikingly like in form to P. arrosor, as 
observed both by Milne-Edwards and Heller, but in place of the scale-like mark- 
ings on the first three pairs of perseopods, here there are quantities of pointed 
tubercles interspersed with fascicles of hairs. 

Locality. — From Station 507, Clarence Bay, Ascension Island, the Scotia obtained 
two specimens, each in a shell of the genus Strombus. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 277 

Genus Eujpagurus, Brandt. 

1851. Eupayurus, Brandt, Middendorff's Sibirische Reise, " Zool.," part i. p. 105. 

1900. „ Stebbing, Pr. Zool. Soc. London, p. 534. 

1905. „ Alcock, Indian Decap. Crust., part ii. fasc. i. p. 122. 

Eupagurus forceps, Milne-Edwards. 
Plate XXVIb. 

1836. Payurus forceps, Milne-Edwards, Ann. Sci. Nat., ser. 2, "Zool.," vol. vi. p. 272, pi. xiii. fig. 5. 

1837. „ „ Milne-Edwards, Hist. Nat. Crust, vol. ii. p. 221. 
1847. Payurus comptus, White, Pr. Zool. Soc. London, p. 122. 

1849. Payurus forceps, Nicolet, in Gay's Hist. Chile, •' Zool.," vol. iii. p. 189. 

1849. Payurus gayi, Nicolet, in Gay's Hist. Chile, " Zool.," vol. iii. p. 190, pi. i. fig. 6. 

1858. Eupayuru* comptus, Stimpson, Pr. Ac. Philad., p. 237 (75). 

1891. ,, ,, Mocquard, in A. Milne-Edwards' Mission du Cap Horn, "Crust.," p. 29. 

1900. ,, ,, Stebbing, Pr. Zool. Soc. London, p. 535. 

1905. Eupayurus forceps, Lagerberg, Schwed. Sudpol.-Evp., vol. v. part vii. p. 2, pi. i. figs. 1-3. 

1910. Paijurus forceps, P. comptus, P. yayi, Rathbun, Pr. U.S. Mus., vol. xxxviii. p. 598. 

Lagerberg gives reasons for identifying E. comptus and its varieties with the 
earlier E. forceps (Milne-Edwards). Nicolet says that his E. gayi is taken along 
with E. forceps on the coasts of Chile, and has the greatest affinity with it, but 
differs by the want of spines on the inner feet and the form of the left cheliped, the 
fingers of which are much more robust. Miss Rathbun, in a List of Species occurring 
from Panama to the Island of Chiloe, gives all the above-mentioned three specific 
names separately, but without discussion of their validity. One of the Scotia speci- 
mens agrees so closely with Nicolet's figure of E. gayi in colouring, size, and features 
that there can be no doubt of its belonging to the species so named by Nicolet, 
but the characters on which he relies do not seem to warrant its separation from 
E. forceps. 

Locality. — Port Stanley, Falkland Islands, January 20, 1903 ; depth 2 fathoms ; 
Station 118. 

Eupagurus modicellus, n. sp. 

Plate XXVId. 

Among the numerous species of this genus I have not been able to find one with 

the combination of characters presented by this little form. The front of the carapace 

has a very blunt median projection and the lateral angles not produced. The eye 

peduncles are long, longer than the front. The second antennae have a slender 

acicle, not denticulate, and a long flagellum, strikingly setose, with some of the setae 

standing out very conspicuously. The endopod of the first maxilla is without a 

flagellum, and all the three pairs of maxillipeds have the exopod flagellate. In the 

third joint of the third maxillipeds the inner margin is finely denticulate, with two of 

the teeth larger than the rest. The right cheliped is much the larger. On the inner 

surface of the hand an elevation extends from the base of the finger transversely to 

the base of the wrist, and another from the tip of the thumb curves gently to meet 
TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 9). 39 



278 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

the first at about one-third of its course from the base of the finger, which also on its 
inner side is traversed from tip to base by a curved elevation. On the outer side the 
hand shows a broad, slightly convex surface, bounded by sharp edges, the wrist and 
hand being denticulate on the border which is continuous with the finely serrate, 
strongly curved outer margin of the stout finger. The left chela has slender fingers 
seemingly longer than the palm. The second and third peraeopods have setae on one 
margin and fine spines on the other of the seventh joint, which ends in a pellucid nail. 
The fourth peraeopods are feebly subchelate, the fifth minutely chelate. The pleopods 
are pellucid, very unequally biramose. The telson is unsymmetrical, each lobe edged 
with small spinules. The carapace measured 2 '5 mm. The specific name alludes to 
the extremely modest dimensions of the new species. 

Locality. — Pyramid Point, Ascension Island ; depth 40 fathoms ; Station 507 ; 
June 10, 1904. 

Genus Calcinus, Dana. 

1852. Calcinus, Dana, U.S. JExpl. Exp., vol. xiii. pp. 435, 456. 

1905. ,, Alcock, Indian Decap. Crust., part ii. fasc. i. p. 51. 

1913. ,, Arata Terao, Annot. Zool. Japonenses, vol. viii. part ii. p. 357. 

In this genus the eye-stalks are long and slender, the left cheliped much larger 
than the right, the fourth peraeopod subchelate, the fifth chelate, the pleopods 
biramose, attached to the pleon on the left side of the second to the fifth segments in 
both sexes, the uropods much larger on the left than on the right side. 

Judging by the species represented in the Scotia collection, the first maxillipeds 
are slight in structure, but the second and third pairs of remarkable strength, especi- 
ally in regard to the exopod, the trunk of which in both pairs far exceeds the endopod 
in breadth, and in the second pair reaches far beyond the endopod's fourth joint. 
In the third pair it does not much exceed that joint, but its basal half is much 
broader than the corresponding part in the second pair ; in both the first joint of the 
flagellum is much broader than the following joints, which are armed with the usual 
setae. In the second maxillipeds the third joint of the endopod is much shorter than 
the fourth, but only a little shorter in the third pair. In both pairs the terminal three 
joints are abundantly furnished with setiform spines, very long and crowded in the 
third pair, where this group of joints exceeds in length that of the third and fourth 
joints combined. 

Calcinus talismani, A. Milne-Edwards and Bouvier. 

Plate XXVIa. 

1892. Calcinus talismani, A. Milne-Edwards and Bouvier, Ann. Sci. Nat. Zool, ser. 7, vol. xiii. p. 225. 
1900. ,, ,, A. Milne- Ed wards and Bouvier, Crust. Decap. " Travailleur" and " Talisman," 

p. 173, pi. xxiii. figs. 15-18. 
1905. „ ,, Alcock, Indian Decap. Crust., part ii. fasc. i. p. 164. 

The Scotia specimens agree well with the description and figures given by the 
eminent French authors, except that in one place, by an obvious slip of the pen, they 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 279 

speak of the right cheliped as being the larger, and in the illustrations the two 
chelipeds can scarcely be drawn to a uniform scale, since otherwise they would be far 
from representing the relative proportions. At least in the Scotia examples the 
chelipeds answer to Alcock's generic definition, " the left being vastly the larger." 
The beautiful colouring is unfortunately no longer available in our specimens to 
assist identification. That the ungues are black in the ambulatory legs may be 
common to many species. The slight curvature and basal widening of the long eye- 
stalks may be worthy of mention, and the bareness of the flagellum of the second 
antennae. The length of about an inch corresponds with that assigned to the speci- 
mens obtained at St Vincent by the French expedition. 
Locality. — Porto Grande, shore, St Vincent ; Station 24. 

Tribe GALATHEIDEA. 

1888. Galatheidea, Henderson, Rep. Voy. " Challenger," vol. xxvii. p. 103. 

1913. ,, Doflein and Balss, Ergebn. Deutsch. Tiefsee-Exp., vol. xx. part iii. p. 131. 

Family Galatheid^e. 

1853. Galatheidee, Dana, U.S. Expl. Exp., vol. xiii. p. 1431. 

1910. ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 362. 

1913. ,. Doflein and Balss, Ergebn. Deutsch. Tiefsee-Exp., vol. xx. part iii. p. 131. 

Genus Munida, Leach. 

1820. Munida, Leach, Diet. Sci. Nat., vol. xxviii. p. 52. 

1891. ,, Mocquard, in A. Milne- Edwards' Miss. Cap Horn, "Crust.," p. 32. 

1899. „ G. M. Thomson, Tr. N. Zealand Inst., vol. xxxi. p. 194. 

1905. „ Lagerberg, Schwed. Siidpol.-Exp., vol. v. part vii. p. 6. 

1908. „ Verrill, Tr. Connect. Ac. Sci., vol. xiii. p. 435. 

1909. „ Chilton, Subantarct. Islands of N. Zealand, part xxvi. p. 612. 

1910. „ Rathbun, Pr. U.S. Mus., vol. xxxviii. pp. 559, 601. 

1911. „ Ortmann, Princeton Univ. Patagonian Exp., part vi. p. 659. 

1913. ,, Doflein and Balss, Ergebn. Deutsch. Tiefsee-Exp., vol. xx. part iii. p. 141. 

Munida subrugosus (White). 
1847. Galathea subrugosa, White, List of Crust, in Brit. Mus., p. 66 (without description). 
1852. ,, ,, White, Voy. of "Erebus" and "Terror," pi. iii. fig. 2 (quoted by Dana 

in 1852). 
1852. Munida subrugosal, Dana, U.S. Expl. Exp., vol. xiii. p. 479, pi. xxx. fig. 7, a, b, c. 
1885. „ „ Filhol, Mission de Vile Campbell, " Zool.," p. 425. 

1902. „ „ Hodgson, "Southern Cross," Nat. Hist., p. 232. 

The above selection of references will enable the student to trace the unending or 
at any rate unended controversy which seeks to determine whether Grimothea gre- 
garius (Fabricius) and Munida subrugosus (White) are one and the same or two 
distinct species. Filhol, Mocquard, Lagerberg, with abundant material, affirm 
that alike in old and young specimens the notably expanded last three joints of the 
external maxillipecls distinguish M. gregarius from M. subrugosus, and Ortmann in 
1911, Balss in 1913, agree with them. Additional distinctions are based on the 



280 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

shape of the carapace and on its rostral and antero-lateral spines, which in the 
subrugosus form are more elongate than in gregarius. Dr Chilton, however, in 1909, 
is still unconvinced, and propounds the theory that " the foliaceous maxillipeds of 
Grimothea gregaria are associated with its pelagic habit," and that when it finds a 
deeper dwelling-place successive moults enable it to assume shorter, less foliaceous, and 
more infolded terminal joints to its external maxillipeds. He considers Grimothea 
novsezelandise, Filhol, also a synonym of M. subrugosus. Making up for this loss to 
the genus, Miss Rathbun in 1910 determines that Grimotea gregaria, Guerin, 
1830-1831, is not the G. gregaria (Fabricius), but a distinct species, Munida coheri, 
noticed by Doflein and Balss as M. cocheri. 

The Scotia obtained the cast skin of a large specimen, with well-pronounced 
rostral and antero-lateral spines, which I am satisfied to call M. subrugosus. 

Locality. — Port William, Falkland Islands, January 1903 ; depth 6 fathoms ; 

Station 118. 

Munida gregarius (Fabricius). 

1793. Galathea gregaria, Fabricius, Ent. Syst, vol. ii. p. 473. 

1820. Grimothea gregaria, Leach, Diet. Sci. Nat., vol. xviii. p. 50 (A. Milne-Edwards). 
1891. Munida gregaria, Mocquard, in A. Milne-Edwards' Miss. Cap Horn, "Crust.," p. 32, pi. ii. 
tigs. 1, la-c. 

Without venturing a decisive opinion on the controversy above considered, I can 
affirm that several little specimens obtained by the Scotia belong to the juvenile 
form described and figured by Mocquard as Munida gregaria (Fabricius). The 
fourth joint of the third maxillipeds is not apically produced, and the two following 
joints are distally dilated. 

Locality. — Stanley Harbour, Falkland Islands, surface ; Station 118. 

Tribe HIPPIDEA. 

1849. Hippidea, de Haan, Crust. Japon., decas 7, pp. 200 and xxii. 
1904. ,, Borradaile, Fauna Maldive, vol. ii. part iii. p. 750. 

1910. „ Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 365. 

Family Albuneidje. 

1858. Albunidx, Stimpson, Pr. Ac. Philad., p. 230 (68). 

1878. Albuneidm, Miers, J. Linn. Soc. London, vol. xiv., No. 76, p. 326. 

1893. ,, Stebbing, Hist. Crust., p. 152. 

1907. „ Nobili, Ann. Sci. Nat, "Zool.," ser. 9, vol. iv. p. 142 

Genus Albunea, Fabricius. 

1798. Albunea, Fabricius, Suppl. Ent. Syst, pp. 372, 397. 

1837. „ Milne-Edwards, Hist. Nat. Crust, vol. ii. p. 202. 

1858. Albunxa, Stimpson, Pr. Ac. Philad., p. 230 (68). 

1878. Albunea, Miers, J. Linn. Soc. London, vol. xiv., No. 76, p. 326. 

1893. ,, Henderson, Tr. Linn. Soc. London, vol. v. part. x. p. 409. 

1907. „ Nobili, Ann. Sci. Nat, "Zool.," ser. 9, vol. iv. p. 142. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 281 

Albunea guerinii, Lucas. 

1758. (?) Cancer earabus, Linn., Syst. Nat , ed. 10, vol. i. p. 632 (reprint). 

1849. Albunea symnista, Lucas, Expl. Algerie, "Crust.," p. 27, pi. iii. figs. 2, 2a. 

1853. Albunea guerinii, Lucas, Rev. et Mag. Zool., ser. 2, vol. v. pp. 45-47, pi. i. figs. 9, 9a, b, b, c. 

1858. Albunxa guerini, Stimpson, Pr. Ac. Philad., p. 230 (68). 

J87S. Albunea guerinii, Miers, J. Linn. Soc. London, vol. xiv., No. 76, p. 327. 

In regard to this handsome species, Miers says : " It is possible that this is the 
species described by Linnaeus (Syst. Nat., p. 1052) from the Mediterranean, under 
the name of Cancer- earabus." Considering the locality and the unusually full 
description which Linnaeus had given in the earlier edition of his Systema, it seems 
even highly probable that his C. earabus is the species with which we are here 
concerned. That the flattened eye-stalks, tapering to little black dots, should have 
been mistaken for movable parts of a divided rostrum, is not so very surprising for 
times when observers trusted much to unaided eyesight. From the typical species 
of the genus A. symmysta (Linn.), 1758, the present is distinguished by the terminal 
joint of the third perseopods, which is without the narrow linear lobe conspicuous in 
the type. This deficiency, however, it shares with A. microps, Miers, and A. thur- 
stoni, Henderson, from the Bast Indies. The Scotia specimen has a very small 
rostral tooth in the central concavity of the frontal margin ; there are eight very 
distinct teeth on either side of this margin, the outermost four being the largest, 
and the one next the eyes being larger than the intermediate three. A strong spine 
projects over the front border of the pterygostomian region, and a smaller one arms 
the basal joint of the outer antennae. The three terminal joints of the third maxilli- 
peds are ventrally and laterally carinate. The oval telson is dorsally grooved down 
the centre nearly to the end, the apex being drawn out subacutely. Length of cara- 
pace at centre 17 mm., at longest part 23 mm. ; breadth 23 mm. ; first antennae 48 mm. 
long ; telson 7 mm. long by slightly over 4 mm. broad. 

Locality. — James Bay, St Helena, June 2, 1904 ; Station 499. 

Lucas in his fig. 9, 1853, shows a frontal margin with three little teeth near the 
eye, and outside of these six larger ones, stating in his text that the teeth are " ordi- 
nairement au nombre de neuf," in contrast to A. symmysta, in which they vary 
between eleven and fourteen. The figure of a finger is marked 9b, and likewise the 
figure of two pleon-segments with the uropods. 

Macrura genuina. 
Tribe SCYLLAKIDEA. 

1893. Scyllaridea, Stebbing, Hist. Crust, Internat. Sci. Ser., vol. lxxiv. p. 191. 

Family Scyllaridje. 
Genus Scyllarides, Gill. 
1898. Scyllarides, Gill, Science, new ser., vol. vii. p. 98. 



282 REV. T. R. R, STEBBLNC ON STALK-EYED CRUSTACEA MALACOSTRACA 

Scyllarides elisabethw (Ortmann). 

1897. Scyllarus elisabethx, Ortmann, Zool. Johrb., vol. x. p. 270. 

1908. Scyllarides elisabethx, Stebbing, Ann. S. Afr. Mus., vol. vi. part i. p. 30, pi. xxx. 

Locality. — Off St Helena harbour, between 45 and 55 fathoms ; June 2, 1904. 

Family Palinurid,e. 

1888. Palinuridx, Bate, Rep. Voy. " Challenger," vol. xxiv. p. 74. 

G-enus Jasus, Parker. 
1883. Jasus, Parker, Nature, vol. xxix. p. 190. 

Jasus lalandii (Milne-Edwards). 

1837. Palinurus lalandii, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 293. 

1884. Jasus lalandii, Parker, Tr. New Zeal. Inst, for 1883, p. 304. 

1910. ,, ,, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 374. 

1913. Palinurus (Jasus) lalandii, Gilchrist, J. Linn. Soc. London, vol. xxxii. No. 216, p. 225, plate. 

The Scotia obtained this abundant species at the entrance to Saldanha Bay, 
depth 25 fathoms. Dr Gilchrist has recently given an interesting description and 
figure of a larval stage, for which he coins the new term " naupliosoma." 

Tribe PENtEIDEA. 

1888. Penxidea, Bate, Rep. Voy. " Challenger," vol. xxiv. p. 219. 
Family Pen^eid^e. 

1881. Penxidx, Bate, Ann. Nat. Hist., ser. 5, vol. viii. pp. 171, 173. 
1888. „ Bate, Rep. Voy. " Challenger," vol. xxiv. p. 220. 

During the present century many important contributions have been made to our 
knowledge of this family, especially by Alcock, Bouvier, Kemp, and be Man. 

Genus Gennadas, Bate. 

1881. Gennadas, Bate, Ann. Nat. Hist., ser. 5, vol. viii. pp. 171, 191. 

1888. „ Bate, Rep. Voy. " Challenger," vol. xxiv. pp. 229, 339. 

1895. ,, Faxon, Mem. Mus. Comp. Zool. Harvard, vol. xviii. pp. 204, 207. 

1901. ,, Alcock, Indian Deep-Sea Macrura, p. 45. 

1904. ,, Rathbun, Harriman-Exp., " Decap. Crust.," p. 147. 

1906. „ Rathbun, U.S. Fish. Coram, for 1903, part iii. p. 907. 

1906. „ Bouvier, Bull. Mus. Ocean. Monaco, No. 80, p. 1. 

1908. ,, Bouvier, Camp. Sci. Monaco, vol. xxxiii. p. 24. 

1909. „ Kemp, Pr. Zool. Soc. London, p. 718. 

1910. „ (as distinct from Amalopenxus), Kemp, Fisheries, Ireland, 1908, p. 13. 

1911. ,, de Man, " Siboga" Exp., vol. xxxixA. pp. 5, 15. 
1913. ,, de Man, "Siboga" Exp., vol. xxxixA., pis. i., ii. 

In 1911 de Man enumerates fifteen named and two unnamed species assigned to 
this genus, with their distribution. He also thinks it probable that the species from 
the Bay of Bengal, referred by Alcock to G. parvus, Bate, is distinct from that 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 283 

species. The list comprises two species originally described by S. I. Smith as Amalo- 
penseus elegans (1882) and A. valens (1884), and a third which Smith described, 
also in 1884, as Benthesicymus 1 carinatus. 

In this genus the male petasma or " andricum," where, as in many cases, it has 
been carefully figured, appears to be very serviceable for specific distinction. 

Gennadas hempi, n. sp. 

Plate XXVII. 

In spite of its mutilated condition, the specimen here described seems properly 
separable from other known species. It makes a near approach to G. elegans in the 
outlines of the carapace, in the proportions of the third peraeopod, and in having 
only one pair of spines at apex of the strongly sulcate telson. But there are several 
differences. The second joint of the first antennae is not shorter than the third. 
The scale of the second antennae is produced well beyond the little lateral tooth. 
The curved spines behind the narrow apical process of the endopod in the second 
maxilla are four in number, two much more curved than the others. In the first 
maxilliped the exopod is shorter instead of longer than the endopod. The petasma 
is nearer to that of G. calmani, Kemp, than any other that has been figured, but is 
more truncate ; the inner opposed margins are straight, lined with a multitude of 
little coupling hooks. The first pleon segment is ventrally produced to a sharp point, 
and in G. calmani it is said to carry " a very strong, sharply pointed spine," which 
may mean the same thing. The apex of the telson is truncate, not rounded, with 
five plumose setae between the single pair of spines, as in G. bouvieri, Kemp. 
G. calmani has (whether invariably or not is uncertain) eleven setae between two 
pairs of spines, subject to the variation of a single pair. It agrees with our species 
in the four curved spines of the second maxilla, but differs in the longer exopod of 
the first maxilliped. Both species belong to the group which have the fourth joint 
of the third peraeopod longer than the fifth, and in both the fingers of the chela in 
that limb are subequal to the palm, but in G. calmani the chela is half as long as 
the fifth joint, whereas in the new species it is more than half as long. Other differ- 
ences between these closely connected species will be found in the two pairs of 
antennae. In the first pair G. calmani has the second joint not equal to the third, 
but "fully three-quarters the length" of it; in the second pair the scale is scarcely 
at all produced beyond the lateral tooth. Through the condition of the respective 
specimens many features are not available for comparison. As described and figured 
by Bate for his G. parvus, the second pleopods have at the apex of the peduncle 
on the inner side two membranous leaf-like appendages. In Bate's figure these 
show setules on the outer margin distally. In our species one of these is somewhat 
bean-shaped, without any armature ; the other, lying between it and the endopod, 
is oval, having much of its inner and distal margin fringed with little, somewhat 
curved, spinules. One of the stations at which G. parvus was obtained by the 



284 RE\ r . T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

Cliallenger was in lat. 37° 49' N., long. 166° 37' W., mid ocean, North Pacific, trawled 
from the reputed depth of 3050 fathoms, which, as will be seen, exceeds that assigned 
to the Scotia species, here named out of respect to Mr Stanley Kemp. 

Locality.— -Lat. 39° 48' S., long. 2° 33' E., trawl 2772 fathoms ; Station 468. 

A fragment of another Gennadas was taken at Station 450, lat. 48° S., 9° 50' W., 
in company with Petalidium foliaceus. It belongs not improbably to Gennadas 
parvus, Bate. 

Family Leuciferid,e. 

1837. " Leuciferiens," Milne-Edwards, Hist. Nat. Crust., vol. ii. pp. 451, 467. 

1852. Sergesiidee, Dana, U.S. Expl. Exp., vol. xiii. p. 601; " Luciferidse," pp. 636, 668. 

1905. ,, Stebbing, Gilchrist's Marine Invest., "S.A. Crust.," part iii. p. 80. 

Under the last reference an extended notice is supplied of the bibliography of 
this family, in which the primary genus bore the preoccupied name Lucifer, changed 
to Leucifer by Milne-Edwards. 

Genus Petalidium, Bate. 

1881. Petalidium, Bate, Ann. Nat. Hist., ser. 5, vol. viii. pp. 172, 194. 
1888. ,, Bate, Rep. Voy. " Challenger," vol. xxiv. p. 348. 

1896. „ H. J. Hansen, Pr. Zool. Soc. London, p. 936. 

1903. ,, H. J. Hansen, Pr. Zool. Soc. London, p. 52. 

Hansen accepts two species for this genus, P. foliaceus, Bate, and P. obesus 
(Kroyer), the former having no branchia above the fourth perseopod, but the latter 
having at least a rudimentary one in that position. He considers that Petalidium 
is distinguished from Sergestes chiefly by the structure of the pleurobranchial plumes, 
feebly developed on the second maxilliped, most developed over the first and second 
perseopods, but less so over the third maxilliped and third perseopod. The pleuro- 
branchial plumes in Petalidium, he points out, have a much lower number of rows, 
a much lower number of plates in the rows, with the plates much larger, curved 
upwards, and looking much more independent than those in Sergestes. 

Petalidium foliaceus, Bate. 

Plate XXVIII. 

1881. Petalidium foliaceum, Bate, Ann. Nat. Hist., ser. 5, vol. viii. p. 194. 

1888. „ ,, Bate, Rep. Voy. " Challenger," vol. xxiv. p. 349, pi. lx. 

1896. ,, „ Hansen, Pr. Zool. Soc. London, p. 936. 

1903. ,, ,, Hansen, Pr. Zool. Soc. London, p. 52, pi. xi. fig. 1, a-g. 

Almost all the figures of this rare species had been drawn before I realised its 
identity with that partially described by Bate and Hansen. Attention may be 
called to the agreement in this plate with Hansen's fig. la of the rostrum. Bate 
had already noticed the small tubercle .adjoining the dark pigment of the eye. 
Hansen notes as at least sometimes present another tubercle, exceedingly small, 
lower down, both being represented in his fig. Id, and well seen in the Scotia speci- 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 285 

men. Bate speaks of the first joint of the first antennae as furnished with a short, 
obtusely pointed stylocerite. Hansen's fig. lb shows it, as I do, with a very small 
acute tooth near the middle of the outer margin. The mandibles are not in agree- 
ment with Bate's figure, except in regard to the penultimate joint of the palp, which 
is of substantial proportions, not slender, as the mandibular palp is generally said to 
be in species of Sergestes. The first and second maxillae are not notably different 
from those in the last-named genus. In describing the first maxillipeds, Bate speaks 
of the third or outer branch as free from hairs, which is at any rate not always the 
case, though they might easily be missing by accident. The substantial character 
and strongly spinose armature of the second maxillipeds may be judged from the 
figure. The third, fourth, and fifth joints are subequal in length, while the sixth is 
rather longer than any one of them. The seventh joint is short, blunt, narrow at the 
base, widening towards the middle. 

In the first peraeopods the third and fifth joints are nearly equal in length, and 
similarly the much more elongate fourth and sixth, both which are strongly spined, 
the sixth faintly showing division into about fifteen jointlets. The fifth joint has on 
its inner margin four spines of very conspicuous length, and distally a group of short 
curved spines to antagonise with a similar group at the base of the sixth joint, as 
in the genus Sergestes. The fourth perseopod is long and slender, the fifth slender 
and short. 

The Challenger specimens were all females or not fully adult. It is therefore of 
interest now to have one with the petasma of the male uninjured. Its complicated 
structure will be best understood from the figure, although that omits one of the 
median plates which are in contact at the base, and only hints at an additional 
pair lying beneath the slender terminal pair and for the most part concealed by it. 
The numerous hooks with which the various projecting lobes are studded probably 
resemble those which Professor S. I. Smith has figured for his Sergestes robustus, 
enlarging them one hundred diameters (U.S. Rep. Comm. Fish, 1884, pi. viii. 
fig. 6a, 66). The second pleopod of a specimen which I take to be a female has, in 
attachment to the short inner branch, a narrowly laminar appendage of considerable 
length, distally furnished with setae on the surface and apically with several 
unequal spines. The inner branch of the uropods is considerably longer than the 
telson, but shorter than the outer branch. In every case, as with the scale of the 
second antennae, this outer branch is apically damaged, the existing portion having 
the outer margin smooth and unarmed. The telson in the basal half is more or less 
parallel-sided, the distal half strongly tapering and ending abruptly in a sharp point, 
the distal part setose at the sides. 

Length of specimen figured 42'5 mm., of which the carapace occupied 11 '5 mm. 
In lateral view the hind margin of the carapace forms a double curve. Among the 
pleon segments the shortness of the fifth and length of the sixth are conspicuous. 

Locality. — Lat. 48° 00' S., long. 9° 50' W. ; depth 1332 fathoms; Station 450. 

TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 9). 40 



286 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

Tribe CARIDEA. 

1852. Caridea (part), Dana, U.S. Expl. Exp., vol. xiii. p. 528. 

Family Pal^emonid^e. 

1819. Palxmonidx (part), Leach, in Samouelle's Entomologist's Compendium, p. 96. 

1852. Palsejiioninse (subfam.), Dana, U.S. Expl. Exp., vol. xiii. p. 569. 

1888. Palsemonidiv, Bate, Rep. Voy. " Challenger," vol. xxiv. pp. 711, 778. 

1890. ,, Ortniaun, Zool. Jahrb., vol. v. p. 512. 

1894. ,, Alcock and Anderson, J. Asiat. Soe. Bengal, vol. lxiii. part 2, No. 3, p. 17. 

1897. ,, Ortmann, Revista do Mnseo Paidista, N. 2, p. 186. 

1901. „ Rathbun, U.S. Fish. Coram, for 1900, vol. ii. p. 123. 

1902. ,, de Man, Abliandl. Senckenh. Gesellsch., vol. xxv. part iii. p. 763. 

1906. ,, Kemp, Fisheries, Ireland, 1905, p. 127. 

1907. ., Borradaile, Ann. Nat. Hist., ser. 7, vol. xix. p. 472. 
1912. ,, de Man, Ann. Soc. Zool. Be/gique, vol. xlvi. p. 197. 

Those who study the authorities above cited will find that different views have 
been and to some extent still are held as to the proper contents of this family. 
The material, however, on this occasion at my disposal does not require, and would 
not facilitate, any thorough discussion of the subject. 

Genus Leander, Desmarest. 

1849. Leander, Desmarest, Ann. Soc. Entom. France, ser. 2, vol. vii. pp. 87, 91. 

1860. „ Stimpson, Pr. Ac. Sci. Philad., vol. xii. p. 40 (109). 

1888. ,, de Man, Arch. Naturg., vol. liii. p. 559. 

1888. Palsemon, Bate, Rep. Voy. " Challenger," vol. xxiv. p. 781. 

1890. Leander, Ortmann, Zool. Jahrb., vol. v. p. 513. 

1893. ,, Stebbing, Hist. Crust., Internat. Sci. Ser., vol. lxxiv. p. 246. 

1901. Paleemon, Rathbun, U.S. Fish. Comm. for 1900, vol. ii. pp. 123, 125. 

1906. Leander, Kemp, Fisheries, Ireland, 1905, p. 127. 

Leander squilla (Linn.). 

1758. Cancer squilla, Linn., Systema Naturae, ed. 10, p. 632. 

1798. Paleemon squilla, Fabricius, Suppl. Eat. Syst., p. 403. 

1884. Leander squilla, Czerniavsky, Mat. zoogr. Pont, comp., fasc. ii. p. 48 (with synonymy). 

1890. ,, „ Ortmann, Zool. Jahrb., vol. v. p. 522, pi. xxxvii. fig. 15. 

1906. „ „ Kemp, Fisheries, Ireland, 1905, pp. 129, 132, pi. xx. fig. 3, a-e. 

Czerniavsky supplies a vast bibliography and names two varieties. Kemp gives 
a synoptic view of the characters which separate this species from L. serratus 
(Pennant) and L. adspersus (Rathke). In the latter two the palp of the mandible is 
three-jointed, while in L. squilla, as discovered by Calman, it is two-jointed, a 
feature which it shares with L. tenuirostris (Say). The rostrum is armed above with 
from seven to ten teeth, below with three, or rarely two or four, and has a bifid apex. 
The telson agrees with that described later on for L. affinis. In the first antennae 
the shortest fiagellum has the number of free joints subequal to that of the coalesced, 
in the specimen examined the number being in each case eleven, and the two sets 
equal in length. Kemp speaks of the shorter ramus as fused to the longer for about 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 287 

two-fifths of its length. In the second maxillipeds the last joint reaches beyond the 
penultimate without making a neatly continuous curve. The second peraeopods 
reach beyond the scale of the second antennae by the whole length of the sixth joint, 
which is a little longer than the fifth, as the fifth is than the fourth ; the seventh 
joint is rather more than half as long as the palm of the sixth. 
Locality. — St. Vincent, Porto Grande ; Station 24. 

Leander affinis (Milne-Edwards). 

1837. Palemon affinis, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 391. 

1852. Paleemon affinis, Dana, U.S. Expl. Exp., vol. xiii. p. 584 ; (1855) pi. xxxviii. fig. 5, a-g. 

1888. ,, ,, Bate, Rep. Voy. " Challenger," vol. xxiv. p. 782, pi. cxxviii. fig. 5, a, d, i. 

1888. „ „ Witmer Stone, in Heilprin, Pr. Ac. Sci. Philad., pp. 318, 322. 

1890. Leander affinis, Ortmann, Zool. Jahrb., vol. v. p. 521. 

1893. „ ,, Ortmann, Ergebn. Plankton- Exp., vol. ii., G. b., p. 47. 

1901. Palmmon affinis, Rathbun, U.S. Fish. Coram, for 1900, vol. ii. p. 125. 

1910- Leander squilla, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 386. 

This species was named for its affinity to Leander squilla, rather inappropriately 
as it has proved, since the latter species belongs to the group which has the 
mandibular palp biarticulate, while here it is three-jointed. Moreover, in the first 
antennae the shortest flagellum has the number of free joints greater than that of 
the coalesced joints, which is not the case in L. squilla. Bate observes in regard to 
his Australian specimens, as a remarkable circumstance, " that neither of the flagella 
of this pair of antennae has attached to it any of the membranous cilia so common in 
the order." In our specimens the short flagellum is well provided with the usual 
filaments, but they are short, and might easily escape notice. Dana supplies some 
good figures of the mouth-organs. But the outermost division of the first maxilla is 
represented much shorter than it is in our specimens. It is possible that Dana saw 
it foreshortened. The continuity of curve between the last two joints of the second 
maxillipeds is well shown in Dana's figure. The terminal joint of the third 
maxilliped is represented both by Bate and Dana as relatively shorter than it is in 
our specimens, especially in the larger ones. As to the second peraeopods, Dana gives 
the " fingers much less than half the length of the hand " ; Miss Rathbun gives " palm 
1'5 times as long as the fingers"; Bate allows the chela to be "about half the 
length of the palm " ; his figure, however, is in near agreement with Miss Eathbun's 
estimate. In a species ranging from Porto Rico to New Zealand some amount of 
variation may well be expected. Dana describes the apex of the telson by saying 
" extremity of abdomen very narrow, having three minute spinules, and between 
them two longish setae." Bate, speaking of the telson, says : " On each side within 
the margin are three small spinules, and the distal extremity is fringed with a few 
hairs." In our specimens the telson ends in a spine-like median process, the base of 
which joins the lateral margins by slightly oblique lines, with a spine at each angle, 
and between these spines and the median process are two other spines nearly three 



288 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

times as long, separated by two densely plumose setae, nearly as long as the spines 
and emerging from below the median process. The length of the species is given by 
Bate as 24 mm. for the male and 33 mm. for the female ; by Miss Rathbun as 
35 "5 mm. for an ovigerous female; by Dana as two inches for his New Zealand 
specimen ; and though some of the South African specimens do not attain to this size, 
others exceed it, reaching 2f inches. 

Locality. — Saldanha Bay, May 19, 1904, 8-10 fathoms; Reitz Bay, May 20, 1904, 
5 fathoms ; Station 482. 

Leander tenuicornis (Say). 

1818. Palxmon tenuicornis, Say, J. Ac. Sci. Philad., vol. i. part ii. p. 249. 

1837. Palemon natator, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 393 (P. tenuirostre, p. 395). 

1849. Leander erraticus, Desmarest, Ann. Soc. Entom. France, ser. 2, vol. vii. p. 92, text-fig. on p. 93. 

1860. Leander natator, Stimpson, Pr. Ac. Sci. Philad., vol. xii. p. 40 (109). 

1879. Leander tenuicornis, S. I. Smith, Tr. Connect. Aci., vol. v. p. 122. 

1884. Palxmon tenuirostris, Cams, Prodr. Faunx Mediterranex, part ii. p. 474. 

1888. Leander natator, de Man, Arch. Naturg., vol. liii. p. 563. 

1888. Palxmon natator, Bate, Rep. Voy. " Challenger," vol. xxiv. p. 784, pi. cxxviii. figs. 6, 7. 

1890. Leander natator, Ortmann, Zool. Jahrb., vol. v. p. 525. 

1893. Leander tenuicornis, Ortmann, Ergebn. Plankton- Exp., vol. ii., G. b., p. 48. 

1908. „ „ Verrill, Tr. Connect. Aci. Sci., vol. xiii. pp. 326, 377. 

Verrill mentions this species along with Latreutes fucorum and the crabs Lupa 

sayi, Gibbes, and Planes minutus (Linn.), as constant tenants of the Gulf weed. 

Ortmann agrees with de Man in regarding de Haan's Palaemon latirostris as a 

synonym of this species, although de Haan, like Desmarest, describes the lower 

margin of the rostrum as without teeth, while Say gives the rostrum eleven or 

twelve teeth above and six or seven below. A dissected specimen shows twelve 

above and six below ; the teeth in this species are much beset with setae, especially 

on the lower margin. Of the lateral spines of the carapace the lower one stands 

back from the margin, instead of adjoining it as in L. squilla and L. affinis. The 

eyes in several specimens retain traces of two parallel colour bands across the 

ophthalmic area. The first antennae have the shortest flagellum united with its 

partner for a very short space. The mandibles have a very slender two-jointed palp, 

the second joint much the longer. The first maxillae differ very slightly from those 

of the other two species, except that the apical series of spines on the inner plate is 

not continued on to the lateral margin. In the second maxillipeds the terminal 

joint is produced well beyond the penultimate, with no continuity of curve between 

them. In the dissected specimen the first peraeopods have the fourth joint longer 

than the fifth, which is slightly shorter than the sixth ; the fingers of the chela are 

considerably longer than the palm. In the second peraeopods also the fourth joint is 

a little longer than the fifth, which is decidedly shorter than the sixth, but here the 

fingers are only a little longer than the somewhat inflated palm. The telson ends apic- 

ally much as in the other species, except that the long spines are at least five times 

as long as those at the angles, and, though separated by the median process, beyond it 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 289 

they come close together, covering the median setse. Among our specimens one had 
the second perseopods of unequal size, in one both first and second perseopods were 
represented only by blunt-ended, few-jointed stalks, in some cases only one of the 
second pair was present. It is likely that these powerful grasping limbs are much 
subject to injury. The length of an ovigerous female was 40 mm. between apices of 
rostrum and telson. 

Localities— Gulf weed, June 29, 1904, lat. 29° 54' N., long. 34° 10' W. ; June 30, 
1904, lat. 32° ll' N., long. 34° 10' W. ; and July 1, 1904, lat. 33° 53' N., long. 32° 
27' W. ; Stations 537, 538, 539. 

Family Hippolytid^. 

1888. Hippolytidx, Bate, Rep. Voy. " Challenger," vol. xxiv. pp. 503, 576. 

1893. „ Stebbing, Hist. Crust., Internat. Sci. Ser., vol. lxxiv. p. 233. 

1901. „ Rathbun, U.S. Fish Comm. for 1900, vol. ii. p. 113. 

1906. ,, Caiman, Ann. Nat. Hist., ser. 7, vol. xvii. p. 30. 

1907. ,, Borradaile, Ann. Nat. Hist., ser. 7, vol. xix. p. 472. 
1907. „ Caiman, Nat. Antarct. Exp., vol. ii., "Crust.," p. 1. 
1910. „ Kemp, Fisheries, Ireland, 1908, p. 99. 

1912. ,, G. O. Sars, Arch. Naturv., vol. xxxii., Nos. 5, 7, 9. 

Genus Hippolyte, Leach. 

1814. Hippolyte, Leach, Edinb. Encycl, vol. vii. p. 431. 
1860. Virbius, Stimpson, Pr. Ac. Sci. Philad., p. 35 (prodromus, p. 104). 
1878. Caradina, Bate, J. Roy. Institution Cornwall, vol. v. p. 486. 
1888. Hippolyte, Bate, Rep. Voy. " Challenger," vol. xxiv. pp. 576, 587. 

In his Challenger Report Bate recognises that as Hipijolyte was originally 
founded for the single species H. varians, Stimpson, by including that species in 
Virbius, condemned his own genus as a synonym to that named by Leach, and for 
the same reason among others Bate saw that his isolation of a species as Caradina 
varians (Leach) must be withdrawn. At the same time he notices that Milne- 
Edwards' genus is Caridina, not Caradina, but prints Verbius in place of Virbius. 

In this genus the mandible has a cutting edge and molar but no palp. The first 

peraeopods have a short stout .chela, the second have a more slender chela and a 

triarticulate wrist. 

Hippolyte acuminatus Dana. 

1852. Hippolyte acuminatus, Dana, U.S. Expl. Exp., vol. xiii. p. 562 ; (1855) pi. xxxvi. fig. 1, a-e. 

1860. Virbius acuminatus, Stimpson, Pr. Ac. Sci. Philad., vol. xii. p. 36 (105). 

1888. Hippolyte bidentatus, Bate, Rep. Voy. "Challenger," vol. xxiv. p. 591, pi. cv. figs. 1, 2. 

1893. Virbius acuminatus, Ortmann, Ergebn. Plankton- Exp., ii. G. b., p. 46. 

This species has been efficiently described and figured by Spence Bate, who 
named it bidentatus, evidently in allusion to the pair of teeth extending subdorsally 
from the hind margin of the fifth pleon segment. Dana takes no notice of this 
character. But these little pellucid processes are not very easy to observe, and the 
probability is that Dana overlooked them. Though a single tooth on the under side 
of the rostrum appears to be usual, one of the Scotia specimens has three teeth 



290 REV. T. R. R. STEBBING ON STALK- EYED CRUSTACEA MALACOSTRACA 

in that position. Bate says that the stouter flagellum of the first antennae is 
" divided into about ten or twelve articuli," which agrees with the Scotia specimen, 
that having eleven joints in one member of the pair and twelve in the other ; but in 
both, contrary to Bate's figure, the two terminal joints are narrow, these and the 
preceding joint being without filaments. Dana describes this flagellum as five- or 
six-jointed, and figures it accordingly. He assigns twelve to fourteen joints to the 
longer flagellum, Bate gives it twelve or fifteen, and our specimen shows seventeen. 
Dana gives the length as varying from three-quarters of an inch to an inch, and this 
agrees with the Scotia specimens. 

Localities.— Gutt weed, lat. 18° 43' N., long. 27° 46' W. ; 27° 23' N., 33° 06' W. ; 
32° 11' N., 34° 10' W. ; Stations 532, 536, 538. 

Genus Latreutes, Stimpson. 

1860. Latreutes, Stimpson, Pr. Ac. Sci. Philad., vol. xii. p. 27 (96). 

1888. ,, Bate, Rep. Voy. " Challenger," vol. xxiv. pp. 576, 581. 

1890. „ Ortmann, Zool. Jahrb., vol. v. p. 505. 

1893. „ Stebbing, Hist. Crust., Internat. Sci. Ser., vol. lxxiv. p. 234. 

1893. ,, Ortmann, Ergelm. Plankton-Exp., vol. ii. G. b., p. 47. 

1901. „ Rathbun, U.S. Fish. Comm. for 1900, vol. ii. p. 114. 

1906. ,, Caiman, Ann. Nat. Hist., ser. 7, vol. xvii. pp. 31, 33. 

1907. „ de Man, Tr. Linn. Soc. London, vol. ix. part xi. p. 421. 
1914. ,, Balss, Abhandl. K. Bayer. Ak. Wiss., vol. ii. part x. p. 46. 

Ortmann and Calman include part of Bate's genus Platybema under Latreutes, 
and Bate's Platybema rugosus is transferred by Calman to a new genus Trachycaris 
on the ground that Platybema was nullified through Bate's erroneous supposition 
that de Haan's planirostris was in generic agreement with rugosus. Had he left 
de Haan alone, his genus would have been unimpeachable. According to Ortmann, 
1893, confirmed by Calman, 1906, Stimpson was in error in attributing an epipod to 
each of the first four peraeopods instead of only the first three. It may be noticed 
also that Milne-Edwards, in describing the type species, assigns only two subdivisions 
to the wrist of the second perseopods, whereas there are three. 

Latreutes fucorum (Fabricius). 

1798. Palxmon fucorum, Fabricius, Suppl. Ent. Syst., p. 404. 

1837. Hippolyte ensiferus, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 374. 

1852. ,, ,, Dana, U.S. Expl. Exp., vol. xiii. p. 562. 

ferus, Stimpson, Pr. Ac. Sci. Philad., vol. xii. p. 27 (96). 

Bate, Rep. Voy. " Challenger," vol. xxiv. p. 583, pi. civ. fig. 1, d-i, k-m, q. 

Ortmann, Zool. Jahrb., vol. v. p. 507. 

Ortmann, Ergebn. Plankton- Exp., vol. ii. G. b., pp. 47, 60. 

Kathbun, U.S. Fish. Comm. for 1900, vol. ii. p. 114. 

Caiman, Ann. Nat. Hist., ser. 7, vol. xvii. p. 33. 
1910. Hippolyte ensifera, Stebbing, Ann. S. Afr. Mus., vol. vi. part iv. p. 390. 

Bate says: " Palsemon fucorum, Fabricius (Suppl. Entom. Syst., p. 404), un- 
doubtedly belongs to this genus." He was deterred from identifying it with the 



1860. 


Latreutes ensift 


1888. 


n n 


1890. 


)> 51 


1893. 


•1 1! 


1901. 


)) !) 


1906. 


»» )> 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 291 

present species because Fabricius describes the carapace as smooth, without mention 
of the little tooth on the gastric region. But, at the date when Fabricius wrote, 
such a tooth might easily have escaped notice or not have been thought worth 
mentioning as interfering with the general smoothness of the carapace. On the 
other hand, among the Malacostraca frequenting the Gulf weed there is, I believe, 
no other species with a smooth, apically quinquedentate rostrum such as Fabricius 
assigns to his P. fucorum, so that I cannot agree with Ortmann's verdict in 1893 
that it is a quite apocryphal species. So far as I have observed, the serration of the 
apex has five points more frequently than any other number, but the variation 
extends from one to nine, the only number within these limits that I have not found 
being eight. The antero-lateral corners of the carapace are serrate, according to Bate 
with five or six points, according to Kathbun with five to eight. In one of our 
specimens there were seven points on one corner and eight on the other. Another 
specimen has eleven points on one side ; those on the other side were not counted. 
Fabricius gives the size as only a third of P. squilla, which well suits the present 
identification, as the length of L. ensiferus varies in different estimates from half an 
inch to an inch. Of those taken at Station 539, it was recorded that one was quite 
blue, two others brown and blue. 

Localities.— Gulf weed, Stations 532, 533, 537, 538, 539, between 18° 43' N., 
27° 46' W., and 33° 53' N., 32° 27' W. 

Some specimens were seen to be infested by the minute Bopyrid Bopyrina 
latreuticola (see p. 301). 

Genus Nauticaris, Bate. 

1888. Nauticaris, Bate, Rep. Voy. "Challenger," vol. xxiv. pp. xii, 577, 602. 
1893. ,, Stebbing, Hist. Crust., Internat. Sci. Ser., vol. lxxiv. p. 234. 

1902. Merhippolyte, Hodgson, "Southern Cross" Exp., "Crustacea," p. 233. 
190fi. Nauticaris, Caiman, Ann. Nat. Hist., ser. 7, vol. xvii. p. 30. 

In Calman's synopsis of the Hippolytidx this genus is distinguished as having 
arthrobranchise at the bases of the first four pairs of perseopods, mandibles with palp 
but without cutting edge, wrist or fifth joint of second perseopod with more than 
seven subdivisions, and a movable spine at the base of the uropods. The last of 
these characters seems to be obscurely expressed, since the spine-like process on the 
peduncle of the uropods is not movable, and the movable spine is at the infero-lateral 
hind corner of the sixth pleon segment. This spine is said by Bate to be absent 
from his species N. unirecedens, though he includes it in his account of the genus. 
But de Man (1907) and Calman agree in identifying N~. unirecedens with the earlier 
Hippolysmata vittatus, Stimpson. Dr Calman points out that Hippolyte magellayiicus, 
A. Milne-Edwards, 1891, belongs to Nauticaris, but differs from other species in 
possessing exopods on the third maxillipeds. He finds that Merhippolyte australis, 
Hodgson, 1902, is synonymous with N. marionis, Bate, Hodgson having been misled 
partly by my acceptance of Bate's error as to the second pereeopods and partly by 



292 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

a wrong view of the mandibles. Hodgson had already noticed the similarity between 
his species and that of A. Milne-Edwards'. 

Here, as in various other instances, the armature of the upper and lower margins 
of the rostrum seems to be so variable within each species that it is an unsafe guide 
for specific distinction. 

Nauticaris brucei, n. sp. 

Plate XXIX. 

Along with a general resemblance to Bate's N. marionis, the present species 
exhibits some rather notable differences in detail. In the first two specimens 
examined the lower margin of the rostrum showed four teeth, but a third specimen 
had only three. A fourth specimen had the hindmost tooth on the carapace more 
remote from the succeeding tooth than in other specimens, a character for which 
Bate named the species N~. unirecedens, mentioned above. The frontal margin of 
the carapace in the new species has only the antennal tooth, without that of the 
infero-lateral angle seen in 1ST. marionis and more conspicuously in N. magellanicus. 
The eyes are much broader at the base than those figured by Bate for iV. marionis. 
In the first antennae the stylocerite does not nearly reach the extremity of the first 
joint. The three-jointed palp of the mandible is relatively much smaller than that 
figured for the species compared. The first maxillipeds have the termination of the 
endopod long and distinctly three-jointed, not a short simple process as figured by 
Bate for N. marionis. For his N. australis Hodgson says, " The endopodite is a 
two-jointed appendage with a terminal claw," and, judging by Mrs Sexton's figure, 
this would seem very distinct both from the present species and from Bate's. In 
the second maxillipeds the fourth joint, much shorter than the third, is quite distinct, 
as the muscles sufficiently attest, but probably the differences in Bate's account of 
his species are due to imperfect observation. The third maxillipeds have a slight, 
easily detachable exopod. The slender second peraeopods have fifteen subdivisions 
of the wrist, those at the base feebly indicated. The apex of the telson has a pair of 
plumose setae between two long spines which are flanked by a pair of short ones. 
Length of specimen figured in bent position 15 mm., straightened out over 20 mm. 

Locality. — Gough Island, depth 100 fathoms; Station 461. 

Specific name given out of respect to Dr W. S. Bruce, leader of the Scotia 

Expedition. 

Nauticaris magellanicus (A. Milne-Edwards). 

1891. Hippolyle magellanicus, A. Milne-Edwards, Mission de Cap Horn., "Crust," p. 46, pi. v. 

figs. 2, 2a-i. 
1902. Merhippol gte magellanicus, Hodgson, "Southern Cross" Exp., "Crustacea," p. 235. 
1906. Nauticaris magellanicus, Caiman, Ann. Nat. Hist., ser. 7, vol. xvii. p. 31. 

According to Calm an this species " differs from the other species of the genus in 
possessing exopods on the third maxillipeds." As already observed, a rudimentary 
exopod is found in the small new species N. brucei. Here the exopod, though short, 



OE THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 293 

is many-jointed and furnished with plumose setae. Apart from this it would be 
difficult to separate this form from Hodgson's Merhippolyte australis, which Calman 
has identified with Bate's Nauticaris marionis. 

The teeth continued from the carapace along the upper margin of the rostrum 
are given by Milne-Edwards as six or seven, with one or two below. Our specimen 
has eight above and one below. The stylocerite of the first antennas reaches beyond 
the main body of the first joint. The scale of the second antennae extends beyond 
the rostrum. The three-jointed palp of the mandibles is furnished on all the joints 
with numerous setiform spines, the first joint much wider and longer than the second, 
the second much wider but little longer than the third. In the first maxillipeds the 
last three joints of the endopod are very different from those of N. brucei, the first 
of them broad, fringed on the inner margin with eleven long setae ; the second longer, 
narrower, curved, somewhat similarly fringed ; the third very short and narrow, 
tapering, straight, tipped with a short seta. In the third maxillipeds the exopod 
is about 2 '5 mm. long, while the last joint of the endopod measures about 8 mm. 
On one of the second peraeopods fourteen components of the wrist are found. In the 
spine armature of the seventh joint in the simple legs there are slight differences 
between this species and N. brucei, but the specimens being so different in size, no 
importance can be attached to these. Similarly, it may be doubted whether slight 
variations in the armature of the telson have any specific value, but the larger apical 
pair of spines are relatively much longer in the smaller of the two forms. The cara- 
pace of the present species measures 14 mm. from apex of rostrum to the middle of 
the concave hind margin ; the scale of the second antennae is fully 6 mm. long. 

Locality. — Port Stanley, Falkland Islands ; Station 118. 

Family Pasiph.eid^. 

1852. Pasiphasidx, Dana, U.S. Expl. Exp., vol. xiii. pp. 532, 536. 

1884. Pasiphaidee, S. I. Smith, U.S. Fish. Comm.for 1882, p. 381. 

1888. Pasiphxidee, Bate, Rep. Voy. " Challenger" vol. xxiv. pp. 481, 857. 

1890. ,, Ortmann, Zool. Jahrb., vol. v. p. 463. 

1893. Pasiphaidse, Wood-Mason and Alcock, Ann. Nat. Hist., ser. 6, vol. xi. p. 161. 

1893. Pasiph&idx, Stebbing, Hist. Crust., Inteinat. Sci. Ser., vol. lxxiv. p. 251. 

1895. Pasipheeiidse, Faxon, Mem. Mus. Comp. Zool., vol. xviii. p. 173. 

1901. Pasiphxidse, Alcock, Gatal. Indian Deep-Sea Macrura, pp. 55, 57. 

1902. „ Rathbun, Pr. U.S. Mus., vol. xxiv. p. 904. 
1904. „ Rathbun, Hariiman-Eocp., "Decapods," p. 19. 

1906. „ Rathbun, U.S. Fish. Comm.for 1903, part iii. p. 927. 

1907. ,, Coutiere, Bull. Inst. Oceanogr. Monaco, No. 104, pp. 1, 12 (larval forms). 
1910. ,, Kemp, Fisheries, Ireland, 1908, pp. 35, 36. 

The genera of this family may be briefly distinguished as follows : — ■ 

(Mandibles without palp, 2. 
Mandibles with palp, 3. 
TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 9). 41 



294 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

(Telson acute or truncate. . 1. Pasiphsea, Savigny, 1816. 

2 ' (Telson with bifid apex . . 2. PJiye, Wood-Mason, 1893. 

(Mandibular palp foliaceous . 3. Psathyrocaris, Wood-Mason, 1891. 

' [Mandibular palp slender, 4. 
Fifth perseopods longer than 
fourth ; second maxillipeds 

having an epipod . . 4. Parapasiphae, S. I. Smith, 1884. 

Fifth pereeopods not longer than 
fourth ; second maxillipeds 
without epipod . . 5. Sympasiphsea, Alcock, 1901. 

Of these genera the first two are closely allied, yet Phye, by its incised telson, is 
different from all the rest, this feature being unfamiliar except in the Schizopoda and 
some larval forms. The third genus is so different from the others that, as Alcock 
has suggested, it " might perhaps be separated to form a distinct family." Lepto- 
chela, Stimpson, 1860, agrees with it in having a foliaceous palp to the mandibles, 
but the palp is one-jointed instead of two-jointed, and other features make its inclu- 
sion in the Pasiphseidse open to question. Bate's imperfectly defined Orphania, 
1888, is perhaps not distinct from Pasiphsea. 

Genus Phye, Wood-Mason. 

1893. Phye, Wood-Mason, Ann. Nat. Hist., ser. 6, vol. xi. p. 164. 
1901. Pasiphsea (Phye), Alcock, Catal. Indian Deep-Sea Macrura, p. 61. 

The original definition is: "Differs from Pasiphae in the carapace and abdomen 
being more or less extensively and distinctly carinated dorsally, in the former being 
armed in front with a pair of branchiostegal spines, and in the telson being forked 
at the extremity." It is said to include P. princeps, Smith ; P. acutifrons, Bate ; and 
P. forceps, A. Milne-Edwards, 1891 ; and characters are given distinguishing these 
species from Phye alcocki (Wood-Mason), 1891. In 1901 Alcock treats it as a sub- 
genus, saying: "Differs from Pasiphsea in having the tip of the telson forked. In 
all other respects, including the number and arrangement of the gills, it agrees with 
Pasiphsea." It should be noticed that Pasiphsea truncatus, Rathbun, 1906, and 
P. Jiagellatus, Rathbun, 1906, both have the apex of the telson truncate, while in 
P. kahviensis, Rathbun, of the same date, the telson has " its tip cut in a very 
shallow V." Nothing could be more satisfactory if we are looking for missing links 
or more confusing to the interests of sharp definition between genus and genus. 

Phye scotise, n. sp. 

Plate XXX. 

By the elongation of the carina of the carapace to a smooth-pointed process 
reaching beyond the eyes, this species is distinguished from the other members of the 
genus. The point of the process is upturned, while in the somewhat similar process 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 295 

of Pasiphsea amplidens, Bate, the point curves downward and does not reach 
beyond the eyes. From Bate's specimen part of the peraeon and all the pleon were 
unfortunately missing. In having the fourth joint of the first perseopod smooth and 
that of the second serrate it agrees with our species, but apart from this there are 
several differences. The mandibular cutting edge here shows nine to ten teeth, 
compared with thirteen in P. amplidens. The first maxilla of the latter is said to 
agree closely with that of P. cristatus, Bate, which according to the figure has the 
middle lobe fringed with seven spines and two spinules, against the eighteen spines 
of the Scotia species. Moreover, here the fingers of the second peraeopods are 
fully as long as the palm, and the fourth joint has only seven teeth along the 
margin, while Bate's figure shows fifteen teeth along that margin and fingers much 
shorter than the palm. 

The telson is about five times as long as its greatest breadth, narrowing towards 
the apex and widening a little at the fork, which is occupied by eight pairs of 
graduated spines. In the first antennae the acicle does not reach the end of the first 
joint, which is as long as the second and third combined, the second being consider- 
ably shorter than the third ; of the two flagella, one for a space is much broader than 
the other. The scale of the second antennas is much narrowed at the flattened apex, 
beyond which the terminal tooth is well produced. The first and second maxillipeds, 
as noticed by Bate, are as firmly attached as if they together formed a single 
appendage. 

Localities.— Lat. 71° 22' S., long. 16° 34' W., depth 1410 fathoms; Station 417, 
March 18, 1904. A second smaller specimen, with anterior process broken off, was 
obtained at lat. 68° 32' S., long. 12° 49' W., by the vertical net from surface to 600 
fathoms; Station 422, March 23, 1904. 

Phye rathhunsB, n. sp. 
Plate XXXI. 

In this species the central carina of the carapace is not produced over the eyes, 
and the telson is only about three and a half times as long as the greatest breadth, 
its apical fork fringed with nine pairs of graduated spines, of which the innermost 
pair is minute. In many respects its proportions differ little from those of the 
preceding species, but in the first antennae the flagella are less unequal at the base, 
and in the second the distal tooth of the scale is less prominent, the middle lobe of 
the first maxilla is fringed with thirteen spines, the first peraeopods have the fourth 
joint serrate with nine teeth and the fingers more instead of less than half the 
length of the palm ; in the second peraeopods the second and third joints as well as 
the fourth are serrate, the fourth joint having as many as seventeen teeth or small 
spines, and the fingers are rather shorter than the palm. 

The earlier-known species, with which the present is most closely related, is that 



296 REV. T. R. R. STEBBING ON STALK- BY ED CRUSTACEA MALACOSTRACA 

which Kroyer in 1845 described as Pasijihae tarda, redescribed and figured under 
the same name by Mr Kemp in 1910. Here, however, instead of the first five pleon 
segments and much of the sixth being sharply carinate dorsally, the earlier segments 
show no carina at all and on none is the carina sharply developed. In the fourth 
perseopods the sixth joint is three-quarters the length of the fourth, instead of less 
than half, as in P. tardus, and in the fifth pair the sixth joint is decidedly shorter 
than the fourth, not about equal to it. In some other respects there may be traced 
divergence between the two species, but with only one specimen of the new form 
available, it is inexpedient to lay too much stress on minutiae. The distribution of 
Kroyer's species extends northwards to Davis Straits and the coasts of Greenland, 
but according to Kemp lat. 51° 54' N. is the most southern locality from which the 
species has been recorded in East Atlantic waters, a very different part of the globe 
from that in which the Scotia specimen was obtained. 

Locality— -Lat. 48° 00' S., long. 9° 50' W. ; depth 1332 fathoms; Station 450. 

The specific name is given out of respect to Miss M. J. Rathbtjn, the distinguished 
American carcinologist. 

A species possibly belonging to this family was taken in lat. 39° 48' S., long. 
2° 33' E. ; depth 2645 fathoms ; Station 468. Unluckily it is represented only by 
the pleon, and that wanting the first pleopods. It is worth mentioning, as the 
remaining pleopods by their great length and slenderness make some approach to 
those in the genus Psathyrocaris, although here the exopod is scarcely more than 
twice the length of the endopod. The sixth segment of the pleon is long and clearly 
carinate ; the telson has two dorsal carinse, which are rather wide apart at the base. 
The exopod of the uropods is elongate. 



Family Nematocarcinidse. 

1884. Nematocarcininie, S. L Smith, U.S. Fish Comm. for 1882, p. 368. 
1888. Nematocarcinidse, Bate, Rep. Voy. " Challenger," vol. xxiv. pp. xiii, 481, 927. 
1893. „ Stebbing, Hist. Crust., Internat. Sci. Ser., vol. lxxiv. p. 249. 

1901. ,, Alcock, Catal. Indian Deep-Sea Macrura, pp 56, 85. 

1910. „ Kemp, Fisheries, Ireland, 1908, pp. 35, 75. 

Alcock remarks that " the Nematocarcinidse in a logical system should not be 
separated as a distinct family, for they are merely Pandalidse in which the first four 
pairs of thoracic legs have delicate exopodites, and they might be united with the 
latter family." But as in the Pandalidse " the thoracic legs never have exopodites," 
and the fifth joint of the second pair is subdivided, while the Nematocarcinidse have 
that joint simple, and in the three following pairs an interlocking arrangement 
between the third and fourth joints described by Bate as of " peculiar and unique 
character," the separation of the two families may very well be upheld, 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 297 

Genus Nematocarcinus, A. Milne-Edwards. 

1881. Nematocarcinus, A. Milne-Edwards, Ann. Sci. Nat. "Zoo!.," ser. 6, vol. xi. art. 4, p. 14. 

1882. Eumiersia, S. I. Smith, Bull. Mus. Comp. Zool. Harvard, vol. x. p. 77. 
1884. Nematocarcinus, S. I. Smith, U.S. Fish. Comm.for 1882, p. 368. 

1888. „ Bate, Rep. Voy. " Challenger," vol. xxiv. pp. lxxxvi, 800. 

1901. ,, Alcock, Gatal. Indian Deep-Sea Macrura, p. 86. 

1906. „ Rathbun, U.S. Fish. Comm.for 1903, part iii. p. 926. 

In this widely distributed genus the names of species are numerous, embracing 
N. cursor, A. Milne-Edwards, 1881 ; Eumiersia ensifera, S. I. Smith, 1882; fifteen 
names given by Bate in 1888 ; and N. agassizii, Faxon, 1893. In 1901 Alcock 
suo-o-ests that Bate's productus, tenuipes, and intermedius may be all the same 
species, and all not improbably synonymous with the N. ensifer of S. I. Smith, to 
which Kemp further refers Bate's Stochasmus exilis as at most a variety, Calman 
and Hansen having already shown that the problematic Stochasmus was founded 
on the young of Nematocarcinus. Alcock questions also the validity of Bate's 
N. undulatipes, supposing that it may be a synonym of N. cursor, to which he 
definitely assigns Bate's N. paucidentatus as a variety. The relative size and 
armature of the rostrum, to which Bate has attached so much importance in dis- 
tinguishing his species, are of doubtful value for that purpose, as they vary with the 
age of the specimen and are otherwise inconstant. There is the further inconveni- 
ence that the long pointed rostrum, like the enormously long perseopods and the 
tapering telson, is peculiarly liable to be damaged. The close resemblance among 
all the accepted species makes it probable that the mouth-organs are practically 
alike in all. But in regard to these it is important to follow the excellent account 
and figures given by S. I. Smith in 1882, rather than those of Bate. In 1893 I 
accepted Bate's statement that the palp of the mandibles was two-jointed, neglecting 
Smith's earlier and correct evidence that it is three-jointed. Smith also shows that 
the second maxillipeds are seven- (not six-) jointed, since the dactylus, which Bate 
overlooks, " is articulated obliquely along the distal end of the propodus." This 
feature, in which the dactylus has quite ceased to be dactyliform, helps to link the 
present genus with several others in the tribe Caridea. Smith has noted that in 
the first maxillipeds the last of the three terminal joints of the endopod is very 
minute. It is indeed difficult to distinguish, but the two preceding are rather long. 
Bate's figure consolidates all three into a tolerably short single joint. According to 
Smith's description of the first maxillae of N. ensifer, " the endognath is much shorter 
than the distal lobe of the protognath, and truncated at the extremity, which is 
armed with a stout seta either side and a third one just below the tip." In our 
specimens these maxillae correspond with Smith's description and figure, except that 
the " palp" or terminal joint which he calls the endognath has at the inner corner of 
its truncate apex a long and strong, distally feathered spine, and at the outer corner 
a much slenderer and shorter spine, and on the surface below the apex a raised 



298 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 

process carrying three or four spines. The second maxillae in our specimens fully 
agree with Smith's account, except that the fringing setse of the proximal lobe are 
much longer relatively than those which he figures. The sharply pointed hind 
portion of the scaphognath has, as he observes, on its inner margin setse of 
remarkable length. 

Nematocarcinus lanceopes, Bate. 
1888. Nematocarcinus lanceopes, Bate, Rep. Voy. " Challenger," vol. xxiv. p. 804, pi. cxxxi. 

Plate XXXIIb. 

The species thus named was taken by the Challenger in lat. 60° 52' S. The 
Scotia obtained a number of specimens in lat. 71° 22' S., which agree fairly well 
with Bate's account of the rostrum, except that the lower margin for almost the 
whole length is unmistakably fringed with setules. A single specimen from lat. 39° 
43' S. has the rostrum devoid of setules. But the more or less damaged condition 
of all the specimens makes it difficult to be sure that any two belong to the same 
species or same variety, while, as above observed, the rostrum itself is subject to 
variation. In none of the specimens examined could I find the fifth perseopod 
complete. According to Bate, the scale or scaphocerite of the second antennae is 
nearly as long as the rostrum. His figure represents it as much longer. In our 
specimens it is decidedly shorter. The third maxillipeds do not reach the end of 
the scale, and are outstripped by the fifth joint of the first perseopods, which almost 
reaches that end. The second perseopods extend much beyond the rostrum. The 
telson reaches the end of the uropods. It carries numerous pairs of little dorso- 
lateral spines, and the narrow apex, besides some setules, has two contiguous spines 
flanked by a much larger subapical pair. The specimens agree in size with Bate's 
specimens, as some of them were about 130 mm. long, allowance being made for 
broken off apex of rostrum or telson. 

Localities.— Lat. 71° 22' S., long. 16° 34' W. ; depth 1410 fathoms; Station 417 
One specimen, probably of the same species, from lat. 39° 48' S., long. 2° 33' E., 2772 
fathoms ; Station 468. 

SCHIZOPODA. 

Order MYSIDACEA. 

Suborder LOPHOGASTRIDA. 

Family EucopiiDiE. 

1852. Eucopidx, Dana, U.S. Expl. Exp., vol. xiii. pp. 601, 609. 

1875. C/ialaraspid.i; v. Willemoes Suhm, Tr. Linn. Soc. London, "Zool.," ser. 2, vol. i. part i. p. 39. 

1883. Eucopiidie, G. O. Sars, Forhandl. Vidensk., Christiania, No. 7, p. 9. 

1910. ,, H. J. Hansen, " Siboga" Exp., vol. xxxvii. p. 19. 

The classification here accepted is fully explained in Hansen's valuable treatise, 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 299 

Genus Eucopia, Dana. 

1852. Eucopia, Dana, U.S. Expl. Exp., vol. xiii. pp. 602, 609. 

1875. Chalaraspis, v. Willemoes Suhm, Tr. Linn. Soc. London, " Zool.," ser. 2, vol. i. part i. p. 39. 

1885. Eucopia, Sars, Rep. Voy. "Challenger," vol. xiii. part xxxvii. p. 54. 

1895. ,, Faxon, Mem. Mus. Gomp. Zool. Harvard, vol. xviii. p. 218. 

1906. „ Ortmann, Pr. U.S. Mus., vol. xxxi. p. 53. 

1910. „ Hansen, " Siboga " Exp., vol. xxxvii. p. 19. 

1912. „ Hansen, Mem. Mus. Comp. Zool. Harvard, vol. xxxv., No. 4, p. 187. 

1913. „ Tattersall, Tr. R. Soc. Edinb., vol. xlix. part iv. p. 868. 

Hansen accepts only the following four specific names as thus far valid in this 
genus : E. australis, Dana, 1852 ; E. unguiculatus, Suhm, 1875 ; E. sculpticauda, Faxon, 
1895 ; and E. major, Hansen, 1910. The extraordinary length of the second, third, 
and fourth perseopods, the delicate structure of the whole organism, and the great 
depth from which it is commonly obtained, have combined to make descriptions 
very imperfect through the provoking mutilation of specimens. 



Eucopia sp. 
Plate XXXIIa. 

As the head, first perseopods, and telson are missing, nothing very definite can 
be said about this species. The remaining limbs come very near to those which 
Sars has figured from a young female which he assigns to E. australis, Dana (Rep. 
Voy. " Challenger," " Schizopoda," pi. x. figs. 2, 5, 6, 7). As in our specimen the fifth 
joint of the fourth perseopod is longer than the sixth, it cannot belong either to Suhm's 
species or to Faxon's. For Hansen's E. major these limbs are not described, nor 
are they distinctly figured by Dana. The spiny armature of the sixth and seventh 
joints does not agree precisely with that shown by Sars. On the sixth joint there 
are five large spines, the first much the longest, separated from the next by one 
spinule, each of the others having two intermediate spinules, the last being followed 
to the apex by eight slender spines. The finger is margined with seven spines, the 
first very small, succeeded by three successively longer, the fourth followed by three 
that are smaller and little projecting. The length might be roughly estimated at 
30 mm., fourth perseopod 39 mm. 

Locality.— Lat. 39° 48' S., long. 2° 33' E. ; depth 2772 fathoms ; Station 468. 

Dr Tattersall's report on the Schizopoda of the Scotia has just appeared, 
and I must apologise for having unwittingly interfered with them, but my plate had 
been finished and the above notice written a long time ago. From the Station 
468 above mentioned Dr Tattersall found the genus Eucopia represented 
by " one fragmentary specimen, head end only," which he groups with damaged 
specimens from various other deep-sea southern localities as probably belonging to 
E. australis, Dana, as redefined by Hansen in 1905. 



300 REV. T. R. R. STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 



STOMATOPODA.* 

1817. " Stomapodes" Latreille, Le Rhjne Animal, vol. iii. p. 40. 
1825. Stomapoda, Latreille, Fam. Nat. du Regne Animal, p. 282. 
1843. Stomatopoda, Krauss, Siidafrik. Crust., p. 60. 
1910. ,, Stebbing, Ann. S. A/7: Mus., vol. vi. part iv. p. 404. 

Family Squillid.e. 

1803. " Squillares," Latreille, Hist. Nat. Crust, et Ins., vol. vi. p. 270. 
1880. Squillidx, Miers, Ann. Nat. Hist., ser. 5, vol. v. pp. 1, 108. 

Squilla armatus, Milne-Edwards. 

1837. Squilla armata, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 521. 
1849. „ „ Nicolet, Gay's Hist, de Chile, " Zool," vol. iii. p. 223. 

1891. ,, „ A. Milne-Edwards, Miss. Cap Horn, "Crust.," p. 53, pi. vii. 

1894. „ ,, Bigelow, Pr. U.S. Mus., vol. xvii. p. 515, text-figs. 9, 10. 

1902. ,, ,, Stebbing, Gilchrist's Mar. Invest., " S.A. Crust.," part ii. p. 45. 

The Scotia obtained a specimen from the stomach of the fish Genypterus 
capensis. The nutritive part of the crustacean was being digested, but the chitinous 
sheath still clearly showed the specific characters briefly noted by the elder Milne- 
Edwards, more fully detailed by Nicolet, and again with good figures set forth 
by Alphonse Milne-Edwards. 

Locality. — Off Dassen Island, between False Bay and Saldanha Bay, South Africa ; 
depth from which the fish was taken, between 30 and 40 fathoms. 

• Larval genus Lysioerichthus. 

1886. Lysioerichthus, Brooks, Rep. Voy. " Challenger," vol. xvi. part xlv. pp. 16, 116. 
1895. Lysierichthus, Hansen, Ergelm. Plankton- Exp., G. c, p. 75. 

This genus contains the larval forms of Lysio squilla, Latreille, 1825. 

Lysioerichthus edwardsii (Eydoux and Souleyet). 

Plate XXVb. 

1837. 1 Erichthus acideatus, Milne-Edwards, Hist. Nat. Crust., vol. ii. p. 501, pi. xxviii. fig. 10. 
1841-1852. Erichthus edwardsii, Eydoux and Souleyet, Voy. "La Bonite," "Zool.," vol. i. part ii. 

p. 260, pi. v. figs. 39-54. 
1852. Erichthus palliatus, Dana, U.S. Expl. Exp., vol. xiii. p. 626, pi. xli. figs. 6, a-e. 
1872. Erichfhoidina armata, Claus, Abhandl. k. Gesellsch. Wiss. Gbttingen, vol. xvi. p. 121, figs. 7, 8. 
1895. Lysierichthus edioardsii, Hansen, Ergebn. Plankton- Exp., G. c, p. 75, pi. vii. fig. 4-4e, 5-5c. 

Hansen, to whom the above synonymy is due, identifies these variously named 
larval forms as all belonging to the adult Lysiosquilla glabriusculus (Lamarck), the 
most advanced stage being E. palliatus, Dana, and the least advanced E. armatus, 
Claus. The Scotia specimen shows all the spine-like projections proper to the 

* For other Stomatopoda collected by the Scotia see Tattersall, Tr. R. Soc. Edin., vol. xlix. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 30 1 

carapace and telson of these juvenile forms. It still carries the obliquely upward- 
pointing medio-dorsal spine not far from the hind margin of the carapace, this being 
a spine which has disappeared from the E. palliatus stage. Nearly equal to it are 
the medio-lateral spines, the position of which is noted as distinguishing these 
larvae from those of Lysiosquilla scabricauda (Lamarck). In a strictly balanced 
dorsal view they are not visible. The narrow part of the rostrum is subequal in 
length to the postero-lateral spines, about 4 mm. in each case. The under margin 
of the rostrum has three microscopic spinules, the middle one the largest ; its apical 
portion is finely serrate on both sides. Adjacent to the eyes is a very small tooth 
on each side, behind which the carapace widens considerably. A minute denticle 
points inward from the base of each postero-lateral process, and between these the 
pleon is clear of the carapace from the third segment. The postero-lateral angles of 
the fifth pleon segment are acute, as are those of the telsonic segment, of which 
the hind margin is finely pectinate and fringed with about forty tiny spinules, its 
centre indented ; the sides have each two teeth, which in this specimen are micro- 
scopic. The slender, strongly geniculate first maxilliped is minutely chelate, the 
small thumb having the inner margin distally finely denticulate ; some of the spines 
on the hand are pectinate. 

Locality.— Lut. 19° 59' N., long. 23° 34' W. ; Station 18. 

ISOPODA EPICARIDEA. 

Family Bopyrid^. 
Genus Bopyrina, Kossmann. 

1881. Bopyrina, Kossmann, Zcitschr. Wiss. Zoo/., vol. xxxv. p. 666. 
1900. „ Bonnier, Trav. zool. Wimereux, vol. viii. p. 364. 

Bopyrina latreuticola (G-issler). 

1845. Bopynis squillarum, Goodsir (not Latreille), Ann. Nat. Hid., vol. xv. p. 75. 

1882. Bopyroides latreuticola, Gissler, American Naturalist, vol. xvi. p. 591, text-figs. 6-8 on p. 593. 

1895. „ „ H. J. Hansen, Ergebn. Planldon-Exp., " Isop.," p. 44. 

1900. Bopyrina latreuticola, Bonnier, Trav, zool. Wimereux, vol. viii. p. 370, text-fig. 61, a-c (from 

Gissler). 
1913. Probopyrus latreuticola, Tattersall, Tr. R.S. Edinb., vol. xlix. part iv. p. 391. 

Bonnier says: "This genus is characterised, in the female, by the absence of 
pleural plates on the pleon, the rudimentary state of the pleopods, of which the last 
pairs have disappeared, as also the uropods, which, however, are still visible in the 
male." Of the species here named no males were discovered. The female is less 
than 2 mm. in length, with the protuberance on one side near the end of the peraeon, 
as delineated by Gissler. 

For the locality see under Latreutes fucorum, p. 291. 

TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 9). 42 



302 REV. T. R. R. STEBBTNG ON STALK-EYED CRUSTACEA MALACOSTRACA 



EXPLANATION OF PLATES. 

Plate XXIII. 

CoryrhyncJius algieola, n. sp. 

n.s. Natural size of -specimen figured above in ventral aspect without limbs and in dorsal aspect with first 

and fifth perseopods on the left and all five peraeopods on the right, and garniture of seaweed 

masking the second peraeopod and partly overlapping its neighbour ; setae for the most part 

omitted. 
r.v. Ventral aspect of rostrum with the right eye, the longitudinally folded first antenna, second antenna 

of the right side, and anterior margin of the buccal frame. 
r.d. Dorsal aspect of the rostrum with its hooked setae. 
T. Distal margin of the pleon of the female. 
a.s. First antenna. 
m., m., mx. 1, mx. 2, mxp. 1, 2, 3. The two mandibles; first and second maxillae ; first, second, and third 

maxilliped8. 
prp. 1. Fingers and part of palm of first peraeopod (cheliped). 
prp. 4. Sixth and seventh joints of fourth peraeopod. 

pip. One of the pleopods of the female, with three of the multitudinous eggs adherent. 
All the figures of separate parts are magnified to a uniform scale. 

Plate XXIV. 

Planes minutus (Linn.). 

n.s. Dorsal view of a specimen natural size, with much magnified view of anterior portion below, and below 
this a similarly magnified view of the ventral aspect, excluding the limbs except the right cheliped. 
a.s., a.i. The first and second antennae. 

mx. 1, mx. 2, mxp. 1, 2, 3. First and second maxillae; first, second, and third maxillipeds. 
prp. 5. Last four joints of the fifth perseopod. 
pip. 1, pip. 4. Rami of the first and fourth pleopods. 
The separate parts are drawn to a uniform scale. 

Plate XXVa. 
Hijmenosoma orbicularis, Desmarest. 

n.s. Dorsal view of female specimen, natural size. The other figures are much magnified, to a uniform scale. 

oc, a.s., a.i. Eye, first and second antennae. 

mxp. 1, 2, 3. First, second, and third maxillipeds. 

prj). 5. Last two joints of the fifth peraeopod. 

The first antenna and the fifth peneopod are from a female specimen, the other appendages from a male 
of the same size. 

Plate XXVb. 
Lysioericldlius edwardsii (Eydoux and Souleyet). 
n.s. Line indicating length of the specimen figured in lateral view. 
car. Dorsal view of carapace, with portions of the eyes, first and second antennae, and one of the medio- 

lateral spines projecting. 
pi. Pleon fnmi the third segment, as seen projecting between the postero-lateral spines of the carapace ; 

part of hind margin much more highly magnified. 
a.s. First antenna. 
mxp. 1, 2, 3. First, second, and third maxillipeds, with distal part of first and third much more highly 

magnified. 
/>/■/>. 1. First peraeopod. 
urp. One of the uropods. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 303 

Plate XXVIa. 
Calrinus talismani, A. Milne-Edwards and Bouvier. 
T. The telsori. 

mx. 2, mxp. 1, 2, 3. Second maxilla ; first, second, and third maxillipeds. 
pip. A pleopod. 

All the figures are drawn to a uniform scale. 

Plate XXVIb. 

Eupagurus forceps (Milne-Edwards). 

mx. 2, mxp. 2, 3, pip. Second maxilla ; second and third maxillipeds j a pleopod. 
All the figures on a uniform scale. 

Plate XXVIc. 

Plagusia capensis, de Haan. 

mx. 2, mx. 2 juv. Second maxilla of P. capensis, and to the same scale that of its immature form 
Maredia elegans. 

Plate XXVId. 
Eupagurus modicellus, n. sp. 

car., a.s., ad. Frontal part of carapace, with eye, and first and second antennae. 

T., urp. Telson and uropods. 

mxp. 1, 3. First and third maxillijieds. 

prp. 1 r., pip. 1 I., prp. 2, 4, 5. First pereeopod, showing the right chela in full from the inner side, and 
the same obliquely from the outer side with the two preceding joints ; the left chela from the 
inner side, inner edges of the tightly closed fingers obscure ; seventh joint of second perseopod ; 
fourth and fifth peraeopods. 
All the figures to a uniform scale. 

Plate XXVII. 

Gennadas kempi, n. sp. 
car. Front of carapace flattened out. 
T. Dorsal view of telson. 
oc. Outline of eye. 

a.s., a.i. First and second antennae, the flagella imperfect. 
to., mx. 1, mx. 2, mxp. 1. Mandible, first and second maxillae, first maxilliped. 
prp. 3. Third peragopod. 

pip. 1, pip. 2. First pleopod Avith the petasma, second pleopod. 
urp. Uropod. 

All the parts are drawn to a uniform scale of magnification, with addition of the telson's apex and the 
apex of the endopod of the second maxilla more highly magnified, in the latter case showing the crowded 
group of four curved spines as viewed from upper and under surfaces. 

Plate XXVIII. 

Petalidium foliaceus, Bate. 

n.s. Specimen partially figured above of the natural size. 

car. , T. Front part of carapace in lateral view ; telson in dorsal view. 

oc. One of the eyes. 

a.s., a.i. Basal joint of first antenna, and base of second with the scale imperfect at the apex. 

???.. Mandible, with further enlargement of the cutting edge. 



304 REV. T. R. R. STEBBLNG ON STALK-EYED CRUSTACEA MALACOSTRACA 

mx. 1 , mx. 2, mxp. 2. First and second maxillae and second maxilliped. 

pip. 1, prp. 5. First and fifth peraeopods in part. 

pet. First pleopod of male with the petasma, one of the median plates omitted for simplicity. 

pip. 2. A second pleopod from a different specimen, probably a female. 

Enlargement of parts to a uniform scale, except in partial figure of the mandible. 



Plate XXIX. 

Nauticaris brucei, n. sp. 

n.s. Line indicating natural size of specimen figured in lateral view, slightly inclined to the right. 

r., car. Rostrum and frontal margin of one side of carapace, with two postocular median spines. 

a.s., a.i. First and second antennae, flagellum of second only in part. 

m., mx. 1, mx. 2, mxp. 1, 2. One mandible and distal end of another, first and second maxillae, and first and 

second niaxillipeds. 
mxp. 3, prp. 1, 2. Third maxilliped, first and second peraeopods. 
prp. 1, prp. 5. Terminal portions of the first and fifth perseopods. 
T. Telson, with distal portion more highly magnified, on a uniform scale with the apices of the first and 

fifth peraeopods and all the mouth-organs except the third maxilliped. The whole telson and 

other separate parts are uniformly magnified on a lower scale. The dissections were not taken from 

the specimen figured entire. 

Plate XXX. 

Phye scotix, n. sp. 

n.s. Lateral view of specimen, natural size. 

T. Dorsal view of telson, with further enlargement of its cleft apex. 
a.s. Peduncle of first antenna. 
a.i. Apex of scale of second antenna. 

*»., to., mx. 1. The two mandibles and the first maxilla, with further enlargement of the maxilla's spine- 
margin. 
mx. 2, mxp. 1, 2, 3. Second maxilla and the first, second, and tbird maxillipeds. 
pip. 1, 2, 4, 5. First, second, fourth, and fifth peraeopods, with further enlargement of the fingers of the 

second. 
urp. Apices of the branches of the uropod. 

All the parts are magnified to a uniform scale, with a scale of further enlargement for the parts above 
indicated. 

Plate XXXI. 

Pliye ralhbunce, n. sp. 

n.s. Lateral view of specimen, natural size. 

car. Anterior portion of carapace. 

T. Telson in dorsal view, with further enlargement of its cleft apex. 

a.s., a.i. First antenna, and second to the base of the flagellum. 

Li., m., mx. 1, 2, mxp. 2, 3. Lower lip, mandible, first and second maxillae, second and tbird maxillipeds, 

with further enlargement of mandible, first maxilla, and second maxilliped. 
prp. 1, 2, 3, 4, 5. The five peraeopods. 
pip. 1, urp. First pleopod and a uropod. 

All parts magnified to a uniform scale, except the front of carapace, which corresponds with the further 
enlargements above specified. 



OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 



305 



Plate XXXI I a. 

Eucopia sp. 

prp. 4. Sixth and seventh joints of fourth perseopod. 
prp. 5. Fifth perseopod. 

Plate XXXIIb. 

Nematocarcinus lanceopes, Bate. 

r. Rostrum, the tapering distal portion missing. 

T. Telson. 

a.i. Scale of second antenna. 

mx. 1. First maxilla, with apex much more highly magnified 

mx. 2, mxp. 1, 2, 3. Second maxilla; first, second, and third maxillipeds. 

prp. 1. First peraeopod. 

All parts magnified to a uniform scale, with the exception above mentioned. 



INDEX. 



Achseopsis . 

aculeatus (Erichthus !) 

acuminatns (Hippolyte) 

sestivus (Lanceola) 

affinis (Leamler) . 

Albunea 

Albuneidse . 

algicola (Coryrhynchus), Plate XXIII 

angulatus (Goneplax) 

anomala (Brachyura) 

anomala (Macrura) 

antarcticus (Lithodes) 

armata (Erichthoidina 

armatus (Squilla) . 

arrosor (Pagurus) . 

australis (Eucoj)ia) 

bipes (Nebalia) 

Bopyrina 

Bopyroides . 

Brachyura . 

brucei (Nauticaris), Plate XXIX. 

Calappidse .... 

Calcinus 

calidus (Pagurus) . 

capensis (Plagusia), Plate XX Vic. 

Caridea .... 

Catometopa . 



PAGE 

257 
300 

289 
257 
287 
280 
280 
259 
264 
272 
274 
274 
300 
300 
276 
299 

257 
91, 301 
301 
257 
292 

271 
278 
276 
267 
286 
264 





PAGE 


Chlorodius ...... 


262 


concavus (Macrocoeloma) . . '. 


261 


Coryrhynchus ..... 


259 


Corystidse ...... 


264 


cristatus (Mursia) .... 


272 


cristimanus (Mursia) .... 


272 


Cyclograpsus ..... 


265 


Cyclometopa ..... 


262 


dormia (Dromia) .... 


273 


Dorynchus ...... 


257 


Dromia . . ... 


273 


Dromiidae .... 


272 


Echidnocerus ..... 


275 


Echinocerus ...... 


275 


edwaidsii (Lysioerichthus), Plate XXVb. 


300 


elegans (Latreillia) . . . 


273 


elegans (Marestia) .... 


303 


elisabethse (Scyllarides) 


282 


ensiferus (Hippolyte) .... 


290 


Epicaridea ...... 


301 


Erichthoidina ..... 


300 


Erichthus ...... 


300 


Eucopia sp., Plate XXXIIa. . 


299 


Eucopiidse ...... 


298 


Eupagurus ...... 


277 


Eurypodius ...... 


260 



306 REV. T. R. R, STEBBING ON STALK-EYED CRUSTACEA MALACOSTRACA 





PAGE 




foliaceus (Petalidium), Plate XXVII 


[. . 284 


Leptodius ....... 


forceps (Eupagurus), Plate XXVIn. 


277 


Leptolitbodes ...... 


fucorum (Latreutes) 


290 


Leucifer . . . . . 


furcillatus (Lispognathus) 


. 258 


Leuciferidse ...... 

Lispognatbus ...... 


Galatheidse ..... 


. 279 


Lithodes ....... 


Galatheidea'. .... 


. 279 


Litbodidse ....... 


Gecarcinidae .... 


. 267 


Lopbogastrida ...... 


Gecarcinus ..... 


. 268 


Lucifer ....... 


Gennadas . 


282 


Lupa ........ 


genuina (Brachyura) 


. 257 


Lysioericbtbus ...... 


genuina (Macrura) 


281 


Lysiosquilla ....... 


Gnatliochasmus .... 


265 




Goneplacidae .... 


. 264 


Macrocceloma ...... 


Goneplax .... 


264 


Macrura ....... 


granulosus (Paralomis) . 


. 276 


maculatus (Grapsus) . . . . . 


Grapsidae ... 


265 


magellanicus (Nauticaris) . . . . 


Grapsus ..... 


265 


Mamaiidae ....... 


gregarius (Munida) 


. 280 


Marestia ....... 


Grimotea ..... 


. 280 


minutus (Planes), Plate XXIV. 


Grimothea ..... 


279 


modicellus (Eupagurus), Plate XXVId. . 


guerinii (Albunea) 


. 281 


Munida ....... 

Mursia ....... 


Halicarcinus 


271 


Mysidacea ....... 


heterochir (Pilumnoplax) 


. 265 




heterochir (Pilumnus) . 


. 265 


naupliosoma ...... 


Heteromacrura 


274 


Nauticaris ....... 


Hippidea 


280 


Nautilocorystes ...... 


Hippolyte .... 


289 


Nebalia ....... 


Hippolytidae .... 


289 


Nernatocarcinidae ...... 


Hymenosoma .... 


269 


Nematocarcinus ...... 


Hymenosomatidae 


. 269 


ocellatus (Nautilocorystes) .' 


Tnachidse .... 


257 


orbicularis (Hymenosoma), Plate XXVa. 


Isopoda ..... 


. 301 


Oxyrrhyncha ...... 

Oxystomata ...... 


Jasus ...... 


282 


Paguridae ....... 


kempi (Gennadas), Plate XXVII. . 


283 


Paguridea ....... 

Pagurus ....... 


lagostoraa (Gecarcinus) . 


. 268 


Palaemonidae . . . . 


lalandii (Jasus) 


282 


Palinuridae ....... 


Lambrus 


. 261 


palliatus (Erich thus) ... 


Lanceola ..... 


257 


Paralomis ....... 


lanceopes (Nematocarcinus), Plate X 


XXIIb. . 298 


Parapasiphae ...... 


latens (Pseudodromia) . 


273 


Parthenopidae ...... 


Latreillia . 


. 273 


parvulus (Xantbodius) . . . . 


latreillii (Eurypodius) . 


260 


parvus 1 (Gennadas) . . . . . 


Latreilliidse . 


273 


Pasiphaea ....... 


Latreutes 


290 


Pasiphaeidae . . . . . 


latreuticola (Bopyrina) . 


291, 301 


Penaeidae ....... 


Leander , 


. 286 


Penaeidea .,,,.. f 



OP THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 



307 





PAGE 




PAGE 


Percnon ...... 


. 267 


Scyllaridse .... 


. 281 


Petalidium . . . 


284 


Scyllaridea .... 


. 281 


Phye .... 


294 


Scyllarides .... 


281 


Pilumnoplax ... 


264 


Sergestes 


. 284 


Plagusia .... 


266 


Sergestidae . 


. 284 


planatus (Halicarcinus) 


271 


Squilla .... 


. 300 


Planes ... 


266 


sqnilla (Leander) 


. 286 


planissimus (Percnon) . 


. 267 


Squillidae 


. 300 


Podochela .... 


. 259 


Stomatopoda . . 


. 300 


Podonema . 


259 


subrugosus (Munida) 


279 


Portunidse . 


263 


Sympasiphaea 


. 294 


Probopyrus . 


301 






Psathyrocaris 


294 


talismani (Calcinus), Plate XXVIa. . . 278 


Pseudodromia 


. 273 


tenuicornis (Leander) . 


. 288 


punctatus (Cyclograpsus) 


265 


thomsoni (Auhseopsis) . 


258 






thomsoni (Dorynchus) . 


258 


rathbunae (Phye), Plate XXXI. 


. 295 










verrucosus (Lambrus) . 


. 261 


sayi (Lupa) ...... 


. 263 






Scliizopoda ...... 


. 298 


Xanthidee .... 


262 


scotise (Phye), Plate XXX. . 


. 294 


Xanthodius . 


262 



Addition to Notice of Mursia cristimanus (p. 272). 

The third, finely illustrated, edition of Cuvier's Regne Animal has two undated 
volumes on Crustacea. Each is entitled " Les Crustaces. Avec un Atlas, par M. 
Milne-Edwards," with the additional word " Texte " in one volume, and " Atlas " in the 
other. Internal evidence makes it completely certain that the text is by Latreille, 
who died in 1833. But it is also clear that the text is independent of the plates, 
since it mentions the genus Mursia (p. 54) without attaching to it any specific name, 
and on p. 262 names Caligus risculus, Leach, for pi. 77, figs. 1, 2, 3, figures which 
Milne-Edwards in his explanation of the plate distributes among three separate 
species of the genus. There may be evidence that pi. 13 of the atlas which records 
" Mursica cristata, Latr.," was published earlier than 1837, the year in which Milne- 
Edwards gives a reference to it under the name " Mursia cristiata," but the atlas 
itself is silent on the point.— T. R. R. S., March 27, 1914. 



nslloy. Soc. Edin r . 



Vol. L.Plate XXIII. 



Stebbing: Stalk-eyed Crustacea Malacostraca. 




R. R. Stebbing 



CORYRHYNGHUS ALGICOLA, n. sp. 



A.RITCH1E & SON. EDIT* »* 



Soc. Edin r . 



Vol. L.Plate XXIV 



Stebbing: Stalk-eyed Crustacea Aalacostraca. 




i. R. Stebbing. 



PLANES MINUTUS (Linn). 



A RITCHIE «. SOU. EDIH 1 ! 1 



Roy. Soc. Edm r Vol. L. Plate XXV. 

Stebbing: Stalk-eyed Crustacea AValacostraca. 




^tt***" ' 



.^-rvT"' > f 



A.— HYMENOSOMA ORBICULARIS, Desnjarest. B.— LYSIOERICHTHUS EDWARDSI1 (Eyd. & Soul.). 



RITCHIE iSOH.EDIN* 



>ar Roy. Soc. Edm r , 



Vol.L. Plate XXVI. 



Stebbing: Stalk-eyed Crustacea /Aalacostraca. 




R. R. Stebbing 

A.— CALCINUS TALISMAN!, A. M.-Edw. & Bouvier. B.— EUPAGURUS FORCEPS (H. M. Edwards). 

C— PLAGUSIA CAPENSIS, de Haan. D.— EUPAGURUS MODICELLUS, n. sp. 



k RITCHIE «. SON. EDIH* 



s.l)y. Soc. Edm r , 



Stebbing: Stalk-eyed Crustacea /Aalacostraca. 



Vol.L. Plate XXVII 




-A H.TCai£ASON,EDW« 



GENNADAS KEMPI, n. sp. 



Is. Roy. Soc. Edm T 



Vol.L. Plate XXVIII. 



Stebbing: Stalk-eyed Crustacea A\alacostraca. 




T. R. R. Stebbing 



PETALIDIUM FOLIACEUS, Bate. 



IE a.SOS.EDIB! 1 



is toy. Soc. Edin r , 



Vol. L.Plate XXIX. 



Stebbing: Stalk-eyed Crustacea /Aalacostraca. 




R. Stebbing 



NAUTICARIS BRUCEI, II. sp. 



Tp 



s Roy. Soc. Edm r . 



Vol. L.Plate XXX 



Stebbing: Stalk-eyed Crustacea /Aalacostraca. 




lT. r. r. Stebbing. 



PHYE SCOTIAE, n. sp. 



A. RITCHIE *.SOW.EDIN* 



y. Soc. Edm r , 



Vol. L.Plate XXXI. 



Stebbing: Stalk-eyed Crustacea Malacostraca. 




R. Stebbing. 



PHYE RATHBUN/E, n. sp. 



(.RITCHIE & SON. BOTH*- 



.s J*/ - . Soc. Edm T 



Vol. L.Plate XXXII. 



Stebbing: Stalk-eyed Crustacea Malacostraca. 




Stebbing. 



A.— EUCOPIA, sp. B. — NEMATOCARCINUS LANCEOPES, Bate. 



( 309 ) 



X. — The Aborigines of Tasmania. Part III. The Hair of the Head compared with 
that of other Ulotrichi and with Australians and Polynesians. By Principal 
Sir William Turner, K.C.B., D.C.L., F.K.S., Knight of the Royal Prussian 
Order Pour le Merite, Emeritus Professor of Anatomy. (With Figures in Text.) 

(Read March 2, 1914. MS. received March 4, 1914. Issued separately June 30, 1914.) 

CONTENTS. 



I'AQE 

Introduction 309 

Tasmanians 310, 334, 340 

New Hebrides 313, 335 

Solomon Islands 317, 336 

New Guinea 317, 336 

Geelvink Bay 317 



PAGE 

Australians 325, 332, 338 

Polynesians 327 

Maoris 327, 338 

Easter Islanders 329, 340 

Hair of Scalp, its Implantation, Form and 

Structure 329 



Papuan Gulf 319 Implantation 330 



Negros of West Africa . . . 319,330,332,336 

South Africa 321 

Bushmen 321, 337, 341 

Hottentots 322, 337 

Kaffirs 323, 337 

Negritos .323 

Andaman Islanders . . . 323, 337, 345 
Semang 324, 338 



Form and Structure 333 

Summary 340 

Melanesians ........ 343 

Bibliography 346 

Explanation of Figures 347 



A number of years ago I began to form and arrange in the Anatomical Museum 
of the University of Edinburgh a collection of the hair of the head to illustrate the 
varieties in colour and character which exist in the Races of Men. In a classifica- 
tion of the races based on the colour and characters of the hair, anthropologists 
have usually adopted the suggestion made by Bory de St Vincent, and have divided 
them into two groups : Leiotrichi, with straight, smooth hair ; and Ulotrichi, with 
woolly or frizzly hair. Each of these again is capable of subdivision. 

In this memoir I intend especially to examine the Ulotrichi, which comprise 

two well-marked subdivisions. In one the hair is very short, and is arranged in 

small spiral tufts, the individual hairs in which are twisted on each other, a mat-like 

arrangement of compact spiral locks closely set together being the result. In the 

other the hair is moderately long, the locks are slender, curled or spirally twisted 

in a part of their length and terminate at the free end in a frizzly bush-like 

arrangement. Ulotrichous hair is found in various African races, in the aborigines 

of Tasmania, New Guinea, the Melanesian Islands in the Pacific, in the Negritos 

of the Malay Peninsula and of some of the islands of the Asiatic Archipelago. The 

Leiotrichi are Australians, Polynesians, Mongols, Malays, Indians, Arabs, Esquimaux 

and Europeans. 

TRANS. ROY. SOC. EDIN, VOL. L. PART II. (NO. 10). 43 



310 PRINCIPAL SIR WILLIAM TURNER ON 

Tasmanians. 

The early voyagers Crozet and Captain Cook, and the French and British 
navigators and naturalists who followed them in the 18th and early years of the 
19th century, as well as the European residents in Tasmania in later years who saw 
the last of the surviving aborigines, described the hair as black, frizzled or woolly, 
usually in short locks, though sometimes forming separate slender ringlets ; with 
abundant beard and whiskers. The statements of these observers have been 
summarised by H. Ling Roth in his comprehensive work on the Aborigines of 
Tasmania,* and in my memoir on these people. 1" This memoir attracted the 
attention of Mr Ling Roth, who with great courtesy and liberality forwarded to 
me in 1908 specimens of the hair of the head of Tasmanians in his possession ; 
also hair of aboriginal Australians from Queensland for purposes of comparison. 
As the hair had been cut from the scalp, its full length and the mode of implanta- 
tion could not be determined. The specimens were accompanied by letters or 
certificates of identity, and permission was granted to describe their characters 
and to add them to the collection in the Anatomical Museum of the University. 

Mr Ling Roth's specimens were as follows : — 

No. 1.- — Hair from the head of a Tasmanian aboriginal chief. This specimen 
was presented in 1898 to Mr H. Ling Roth by Mr James B. Walker of Hobart, 
with the following history : — It was cut from the head of one of the chiefs who 
accompanied G. A. Robinson on his " pacific mission." They visited Mr Pike's house 
at Park Farm, Jericho, sometime about 1832 or later, and Mr Pike's daughter cut 
a lock from the chiefs head. The specimen consisted of a few loose hairs, the 
longest of which measured 4 cm.; each of which was curled and black in colour. 
They are stated to have been pulled out of a lock of hair 17 cm. long and 3 mm. 
thick, which formed a spiral curl so compact that long hairs could not be extracted 
from it. The lock was coated with ochre, or a clayey substance. 

No. 2. — Mr Walker also sent later in 1898 a scrap of Truganini's hair, accom- 
panied by a copy of the following certificate : — " I certify that the accompanying 
is a lock of Lallah Rook's hair (the last of the aborigines), presented by her to 
Mr F. Howell. (Signed) J. S. Dandridge, Oyster Cove, 1st April 1872 " (fig. l). 

The original certificate is in the possession of John Macfarlane, who was present 
at Oyster Cove on 1st April 1872, and at whose request the lock of Truganini's 
hair was obtained and the certificate given. The lock consisted of a few short 
hairs from 3 to 4 cm. long ; it formed a close spiral at its middle, but was looser 
at the periphery. The colour was so dark that it might be called black (fig. 2, A). 

No. 3. — A lock of hair labelled "Hair of Mrs Thos. Cochrane Smith, the so-called 
last of the Tasmanians." Mr Ling Roth in Appendix G to his work gave three 

* Halifax, England, 1899. 

t "The Craniology, Racial Affinities and Descent of the Aborigines of Tasmania," Trans. Roy. Soc. Edin., 
part i. vol. xlvi. p. 365, 1908 ; part ii. The Skeleton, vol. xlviii. p. 413, 1910. 



THE ABORIGINES OF TASMANIA. 



311 



figures from photographs of Mrs T. Cochrane Smith, and recognised that, although 
her facial characters partook largely of those of the Tasmanians, she was not a pure 
aboriginal, but was of mixed blood. The lock was a distinct curl but was not 
spirally twisted; its length was 8 to 9 cm., but when stretched it measured 
11 '5 cm. ; its breadth was about 8 mm. The hairs varied in colour from a darkish 
brown to pale brown, and some were greyish. 







Fig. 1. — T;isniauian, Truganini. 



No. 4. — 1 have subsequently obtained from another source, of undoubted authen- 
ticity, some hairs of a male Tasmanian attached to a fragment of the scalp. The 
hairs emerged close together and formed a lock 4 to 5 cm. long. They were distinct 
and wavy for a short distance, but soon aggregated together and became arranged as 
a compact spiral coil 2 cm. long, beyond which they again separated, and were curly 
and frizzly at the free ends (fig. 2, B). A few delicate hairs, about 6 mm. long, 
ended close to the scalp in pointed ends ; they were so short as not to reach the spiral 
portion of the lock. The dried hair was deep brownish black in colour. 

The illustrations in Ling Roth's volume reproduced drawings and photographs 
of aborigines in some of which the hair was short and the curly tufts were not unlike 



312 



PRINCIPAL SIR WILLIAM TUP.NER ON 



the specimen above described.* In others, again, the hair was much longer and 
formed slender compact ringlets which hung pendulous in front of the forehead 





A B 

Fig. 2. — A, lock from Truganini ; B, from No. 4. 



and down the side of the ears and cheeks, t Several museums possess a cast of the 
bust of a Tasmanian man with this character, and the Anatomical Museum of the 
University is indebted to Professor Anderson Stuart for a copy of one4 In it 




Fig. 3. — Tasmanian, from cast of a bust. 



the ringlets were arranged so as to fall from the crown of the head, either 
forwards, backwards or to the sides (fig. 3). They varied in length from 6 to 
20 cm. Those in front concealed the forehead, and the longer ringlets, about 

* See pp. 9, 33, 41, and Appendix Q— Truganini. t See pp. 17, 25 of his work. 

J The cast is apparently a copy of a bust modelled by Dumoutier, Atlas to Voyage of Dumont d'Urville, 1830. 



THE ABORIGINES OF TASMANIA. 



313 



12 cm., reached the eyebrows and the root of the nose; those on the back and 
sides of the head were the longest and concealed the ears, cheeks, back of the 
neck and reached the shoulders. 

New Hebrides. 

Melanesian aborigines with dark-brown or black skins, dolichocephalic heads, and 
with stature in the men of about 5 feet 6 inches inhabit this group of islands. 
Captain Cook visited them in 177 A* He described the men of Mallicolo as having 
long heads, Hat faces, hair mostly black or brown, short and curly. The people of 




Fig. 4. — New Hebrides, Tanna. 

Erromango had the hair crisp and curly and somewhat woolly. He figured the head 
of a man of Tanna, and said the hair was black and brown, tolerably long, very crisp 
and curly, separated into slender locks around which a thin vegetable fibre was 
wound to about an inch from the free ends. The locks were somewhat thicker than 
whip-cord, and looked like a parcel of small strings hanging down from the crown of 
the head to the back and shoulders. Prichard has also figured t from a drawing by 
Captain Erskine a similar arrangement of the hair on a native of Aneityum. 

The Museum is fortunate in possessing several specimens from the natives of the 

* Second Voyage, vol. ii. p. 78, with plate. 

t Natural History of Man, vol. ii. pi. xxxvii. An account of Captain Erskine's voyage is given in his Journal 
of a Cruise among the Western Islands of the Pacific, London, 1853. 



314 



PRINCIPAL SIR WILLIAM TURNER ON 



New Hebrides, which were presented in 1892 by the Reverend James H. Lawrie, for 
many years a missionary at Aneityum.* On the island Efate the hair was worn short 
by both sexes. On Tanna, Aniwa and Aneityum, and to a limited extent on Futuna, 
the men dressed the hair in the manner described by Cook. Years of attentive 
dressing were required to obtain a coiffure such as Cook described. The practice 
is said to have ceased under the influence of the missionaries. The collection con- 
sisted of seven specimens obtained from diiferent persons : — 

No. 1. — Eight distinct locks from a man, the longest of which was 30 cm. (12 
inches). They had evidently been cut close to the scalp, 2 inches from which a 
thin narrow vegetable fibre had been wound tightly around the lock for from 16 to 
17 cm., beyond which the hair was free, either loosely or compactly spiral or frizzly, 




Fig. 5. — New Hebrides. Three locks from No. 1. 



and the free end was 5 to 8 mm. broad (fig. 5). The hair was brownish black, 
and varied somewhat in the depth of its tint. 

No. 2. — Adult male. The hair in proximity to the scalp was matted and beyond 
the tangle seven slender locks about 10 cm. long (4 inches), 4 to 5 mm. broad, 
had been isolated and carefully plaited for about two-thirds of their length, the 
terminal third being spirally curled and frizzled. The tangled hair was dark brown, 
but it was brownish yellow in the lock itself as if artificially discoloured. 

No. 3. — A bundle of hairs from a native of Futuna Island, around which a white 
band had been fastened. A division into locks was indicated though not complete. 
The hair was frizzly and showed loose spirals. In colour it was dark brownish 
black (fig. 7). 

No. 4. — From an adult male; No. 5 from a young man, pet. 18. Both were 
matted and spirally curled but not in definite locks. The colour was brownish 
or black. 

* "The New Hebrideans" in Scottish Geographical Magazine, June 1892, vol. viii. 



THE ABORIGINES OF TASMANIA. 



315 



No. 6. — Adult female, a number of locks, to some extent matted at their scalp 
ends, but capable of being drawn asunder, when their arrangement in loose spiral 




Fig. 6. — New Hebrides, Tanna. Three locks from No. 2. 



curls about 7 cm. long was seen. When stretched the length of the locks was 
about 13 cm. (5 inches). At the sides and free ends the locks were dishevelled 
and frizzly. The colour was dark brown and black. 




Fig. 7.— New Hebrides, Futuna. No. 3. 



No. 7. — Girl, set. about 14. Seven locks, matted at the deep ends, but with well- 
marked loose spiral curls frizzly at the sides and ends. The locks, without being- 
stretched, were from 5 to 7 cm. long, but could be elongated to 9 cm. Colour 
black with brown tint. 

No. 8. — Some years ago Dr George Porter presented to me the coiffure of 



316 PRINCIPAL SIR WILLIAM TURNER ON 

a Kanaka labourer who had worked on a sugar plantation in Queensland. It was 
17 inches in height and had projected upwards as a top-knot from the crown 
of the head.* It consisted of 834 slender locks tied together about the middle 
with a tape to form a bundle. The hairs in each lock, where it had been cut 
from the scalp, were free, but each lock was then tightly wound by a narrow 
band of vegetable fibre for several inches, beyond which the hairs again separated, 
were loosely spiral and frizzly at their free ends. The hair next the scalp and 
that enveloped by the fibre was brownish black in colour, but the frizzly ends 
were auburn-tinted, apparently bleached (fig. 8). The locks closely resembled No. 1 




Fig. 8. — Probably from native of New Hebrides. No. 8. 

in the Revd. James Lawrie's collection from the New Hebrides. In their arrange- 
ment as a top-knot, the coiffure to some extent resembled the one figured by 
Dr Prichard,! said to be from a native of Ombai,} in which, however, the hair 
did not seem to be dressed in separate locks, though frizzly at the free ends. I 
did not learn the name of the island of which the Kanaka labourer was a native, but 
Mr Lawrie stated in his paper that the New Hebrides is a source of supply for the 
Queensland plantations. 

In his account of the people of New Caledonia, an island to the west of the New 
Hebrides, Captain Cook recognised their resemblance to the aborigines of Tanna, 

* Report, British Association, Edinburgh meeting, p. 906, 1892. 

+ Natural History of Man, vol. ii. p. 441, 1855. j Ombai is an island, east of Java and west of Timor. 



THE ABORIGINES OF TASMANIA. 317 

in the colour of the skin as well as in the black colour of the hair and beard. 
The hair, coarse and strong, was sometimes tied up on the crown of the head, whilst 
at others a large lock was worn on each side. It was much frizzled and mop-like, 
and at first looked as if it resembled that of a negro, but it was, he said, really very 
different. In some of the men, however, and in all the women the hair was cut into 
short locks. 

Solomon Islands. 

These islands are occupied by Melanesians. Many years ago Dr J. C. Cox of 
Sydney presented to me two skulls from Rubiana, the cephalic index of which, 72 
and 70 respectively, was dolichocephalic* The scalp with hair had been partially 
retained on one skull. The hair was black and curly, as a rule from 3 to 6 cm. long 
and seldom arranged in locks. On the vertex, however, some short locks had been 
preserved, which projected from 2 to 3 cm. ; they formed loose spirals, the hairs of 
which could be stretched to 5 or 6 cm. A few hairs attached to each other at the 
scalp ends were loosely curled, and the longest when stretched attained the length 
of 23 cm. 

Although it is customary to speak of the hair of the Melanesians as black, it has 
not the depth of tint one sees in Negros, Hottentots, Mongols, Indians, Esquimaux 
and Australians, but has a dull brownish admixture, which is more noticeable when 
the hair is held to the light. Sometimes the hair is of a lighter brown colour, a 
change which is possibly to some extent due to a partial bleaching caused by lime 
and clay with which the natives dress the hair. 

New Guinea. 

The attention of the early voyagers to New Guinea was at once attracted by the 
magnificent mop-like arrangement of the hair in some tribes of Papuans. The 
appearance has been frequently described and figured, more especially in the 
aborigines in the south-east of the Island and on the Papuan Gulf, which together 
form British New Guinea, in whom the coiffure may be 3 feet or more in circum- 
ference. In the Fly River district and in Dutch or Western New Guinea the hair 
is kept much shorter, from 4 to 5 inches, and is plaited into pencil-like cords or 
ridges, t The free ends of the hair whether in mops or pencils are frizzled, from 
which character it is said the name Papuan is derived from a Moluccan word 
signifying frizzled (fig. 9). 

Geelvink Bay, Dutch Neiv Guinea. — The late Dr A. B. Meyer presented to 

* Challenger Report on Human Crania, pp. 93, 96, part xxx., 1884. 

t For recent descriptions of the arrangement of the hair in the people of New Guinea, consult Sir Wm. Mac- 
gregor, British New Guinea, Country and People, London, 1897 ; Chalmers and Gill, Work and Adventure in New 
Guinea, London, 1885 ; Brown, Pioneer Missionary and Explorer, London, 1908 ; Seligmann, The Melanesians of 
New Guinea, Cambridge, 1910 ; Wollaston, Pygmies and Papuans, London, 1912 ; Captain Rawling, Land of the 
New Guinea Pygmies, London, 1913 ; Henry Newton, In Far New Guinea, London, 1914. 

TRANS. ROY. SOC. EDIN., VOL, L. PART II. (NO. 10). 44 



318 



PRINCIPAL SIR WILLIAM TURNER ON 



me in 1902 a specimen which he had obtained in 1873, from a Papua boy, six to seven 
years old, at Rubi, Geelvink Bay. It consisted of two locks, each of which was again 




Fig. 9. — New Guinea, Port Moresby. 




Fig. 10. — Locks, New Guinea, Geelvink Bay. 



divided into two (fig. 10). A lock was 18 cm. long (7 inches) and 3 mm. broad. At 
the scalp end the hairs were curly, but in the lock itself they were twisted into a 



THE ABORIGINES OF TASMANIA. 319 

compact pencil-like bundle, at the free end of which the hairs were again wavy. # 
At the scalp the hairs were brownish black, but they changed to reddish brown in 
the lock itself, t Another specimen consisting of separate hairs was also given by 
Dr Meyer : in it the longest hairs, when straightened, were about 17 cm., and their 
curly and wavy character was distinct. The colour was brown-black. 

Papuan Gulf. — I am indebted to Professor A. C. Haddon for two cuttings of 




Fig. 11. — Lock, Papuan Gulf. 

hair, one of which he took off a dance mask, whilst the other was removed from an 
artificial wig, which had probably formed part of a dance mask. In the latter the 
hairs had been cut very short ; they were loosely matted together and were spirally 
coiled. In the former the hairs were partly matted and curly, but one lock was 
distinct, between 6 and 7 cm. long and curled (fig. ll). Both specimens were 
brownish black in colour. 

Negros. 

The hairs on the head of the African negro and his descendants in America and 
elsewhere are arranged in short, compact, black tufts or locks, about 5 mm. in diameter, 
the hairs in which are spirally coiled. The arrangement is best observed from the 
norma verticalis of the head (fig. 12). Over a large part of the surface the locks 
were closely compacted together like a mat, and the hairs in given areas, as they 
emerged from the scalp, at once proceeded to form the constituent parts of their 
respective tufts. On the sides of the head the tufts were further apart, and the 
surface of the scalp between the locks was hairy, equally with that in superposi- 
tion to which the locks were situated, so that no spots of noticeable size bare 
of hair were seen. Locks of hair, conformable to this character, constitute the 
appearance to which the term woolly was originally applied and is the typical 
arrangement. 

The Museum contains a characteristic negro's head from which the above descrip- 
tion has been written ; also several locks of hair from other individuals which have 

* Meyer, A. B., Mitth. Anthro. Ges., Wien, vol. iv. Nos. 3, 4, 1874. 

t Rich. Neuhauss has written in Zeitsch. fur Etlmologie, 45th Jahrgang, Heft iii., 1913, a paper on the Red Blond 
Hair of the Papuans. He recognises bleaching of the hair by lime, but states that in addition to black pigment 
granules a diffused reddish and yellow substance is present in the hair of Papuans. Blond hairs may sometimes co- 
exist along with black hairs. 



320 



principal sir William turner oN 



been cut close to the scalp. The locks in their natural disposition projected about 
l|- cm. ( T % inch) above the surface of the scalp ; the individual hairs when put on the 




Fig. 12. — Norma verticalis, scalp of Negro. 

stretch were elongated to about 5 cm. (2 inches). The shortness of the lock was due 
to the rapidity of the spiral and its consequent compression. To some extent the 





Fig. 13. — Locks of hair from two Negros. 



free ends of the hairs in adjacent locks may become entangled with each other 
(fig. 13). It should be noted that the arrangement is natural to the race and is 
not produced artificially. 



THE ABORIGINES OF TASMANIA. 



321 



South Africa. 

This part of the continent is occupied by races of dark-skinned, woolly-haired 
aborigines (fig. 14). I am indebted to Dr Ian Raubenheimer, from South Africa, 
a graduate of the University, for a number of specimens of the hair of Bushmen, 
Hottentots and Kaffirs, which had been cut from the heads of living persons. 




Fig. 14.— Bush. 



Bushman.- — No. 1. A black, single compact lock 3'6 cm. long, which stretched to 
5 cm. (2 inches) ; its greatest breadth was only 2 mm. and it had a wavy outline. It 
was composed of hairs closely coiled on each other and spirally arranged (figs. 15, 32). 

No. 2. Bushwoman. — A number of short locks loosely aggregated, which when 





Fig. 15. — Bush, partly matted, partly separate locks. 

drawn asunder measured from 1'5 to 2 cm. in length and 3 to 4 mm. broad. The 
individual hairs were spirally wound in the lock, and the collective arrangement 
formed a mat of hair in which the spiral locks could be recognised (fig. 15). 

No. 3. From an old Bushwoman. A similar mat-like arrangement of short locks, 
the constituent parts of which were formed of spirally coiled hairs, whose free ends 
were to some extent intertwined. The colour was greyish brown, obviously from 
senile changes. 



322 



PRINCIPAL SIR WILLIAM TURNER ON 



The hair in each specimen was distinctly woolly, though longer in the man than 
in the woman, and the individuality of the lock was more distinct. The locks were 
recognisably smaller than in the negro. 

Hottentots. — Hair from four Hottentot women. Each specimen had been dressed 







Fig. 16. — Hottentot with plaited locks. 

in artificially three-plaited locks. They varied in length from 6 cm. (2"3 inches) to 
30 cm. (llf inches). The broadest lock was dishevelled at the end cut from the scalp, 
where it was 3 cm. broad ; this was succeeded by the plaited part 5 '8 cm. long, which 




Fig. 17.— Zulu Kaffir. 

at first was 1*5 cm. broad, but at the free end, where the plaiting ceased, was only 
8 mm. broad (fig. 1 6). The individual hairs were curly but not definitely spiral. 
The colour was rich black in all these specimens, except in one which was dark 
brown in colour, slender in its whole length and about 4 mm. broad. The longest 
lock was of almost equal breadth, 7 mm., throughout its entire length. 



THE ABORIGINES OF TASMANIA. 323 

In addition to these specimens were some tufts of hair from children, which con- 
sisted of short, spirally curled locks about 25 mm. long, but capable of being stretched 
to about 35 mm. Their free ends were to some extent intermingled with each other. 
The colour was deep brown, almost black. 

A cutting of hair marked " Bastard Hottentot," from a boy, set. 3, was included 
in the collection. It was not formed of curly spiral locks, but consisted of almost 
straight hairs 3 to 4 cm. Ions and dark brown in colour. The Bastards are the 
children of mixed black and white parents. 

Kaffirs. — A specimen from a woman, set. 35. It consisted of fine, short, com- 
pact, spirally curled locks, the hairs of which were to some extent intermingled so 
as to form a loosely aggregated mat. The locks were black and more slender than 
in the adult negro (fig. 18). Three specimens of the hair of children from ten months 




Fig. 18. — Kaffir locks of hair. 

to three years of age were also received ; their character and arrangement closely 
corresponded to that of the woman. 

The Kaffir hair from its short, spirally curled locks and their close arrangement 
should without doubt be classed as woolly (fig. 17). The Hottentots, on the other 
hand, had relatively long hair, capable of being artificially plaited into ornamental locks, 
an example of which as above recorded was about 12 inches long (fig. 16). They are 
not regarded as a pure race. 

Negritos. 

Pygmy or dwarf races, shy in their habits, living in small communities and dis- 
tinguished by short stature, black or brown-black skins, woolly hair, brachycephalic 
or mesaticephalic skulls, are met with in some tropical countries. 

Andaman Islanders. — Surgeon Brander, medical officer at Port Blair, stated * 
that they shaved the head, the operation being performed by women, who used their 
milk as a shaving soap. Locks of hair are therefore somewhat difficult to obtain, 
and the Museum possesses only two specimens of brownish-black hair, the longer 

* Proc. Roy. Soc. Edin., vol. x. p. 415, 1880. 



324 PRINCIPAL SIR WILLIAM TURNER ON 

of which is 5 "3 cm., though it can be stretched to 9 cm. (3|- inches) ; its average 
breadth is about 3 mm. (fig. 19). Each lock consisted of a compact spiral bundle of 
hairs, which disengaged themselves at the sides and free ends, aud assumed the form 
of short narrow ringlets. I owe to the late Dr Joseph Dougall, who had gained 
the confidence of the natives, a photograph of a group of thirty islanders of both 
sexes, in some of whom the hair was present, though in others the scalp was bare 




Fig. 19. — Andaman Islands locks of hair. 

as if shorn. The dwarf stature is well seen in contrast with that of Dr Dougall, 
who was a tall man, standing in the group (fig. 34). 

Semang. — A tribe living in the interior of the Malay Peninsula described by 
Nelson Annandale and H. C. Robinson.* The Museum is indebted to the former 
for a lock of hair of a member of the Hami tribe, Jalor. It was black in colour, 
woolly, only 2 '2 cm. long, but could when stretched be somewhat elongated. The 
individual hairs were spirally twisted (fig. 20). 




Fig. 20. —Semang locks of hair. 

The Museum does not contain specimens of the hair of the Aetas of the Philippine 
Islands or of the pygmies in Central Africa. The question of the presence of Negrito 
pygmies in New Guinea, which long formed a subject of discussion, has now been 
definitely settled. In 1910 Captain Rawling and Mr A. F. R. Wollaston dis- 
covered and described in their respective volumes of travel a tribe of pygmies in 
the Mimika River district at Tapiro, near the Nassau Mountains in Dutch New 
Guinea,! with short, black woolly hair, black whiskers and beard, chocolate-coloured 

* Fasciculi Malayensis, "Anthropology," part i., 1903. 

t See their books on the New Guinea Pygmies and Papuans, op. cit., p. 317, footnote, and Bibliography. 



THE ABORIGINES OF TASMANIA. 325 

skin, wide nostrils, thick lips, brachycephalic heads, and average stature in the men 
4 feet 9 inches. 

At the same time a Dutch expedition, engaged in exploring the North-West River 
district, situated east of the Mimika River, met with a similar tribe of pygmies in a 
village on a slope of the Goliath mountain. A careful description of the people was 
written by A. C. de Kock, surgeon to the expedition, which has added to our know- 
ledge of the New Guinea pygmies.* A. J. P. v. d. Broek in a recent article t has 
compared de Kock's written descriptions with those of the English explorers, and 
has reproduced photographs of the people, whose physical characters strictly conform 
to those of other pygmy races. What is especially interesting in connection with 
this memoir is a description and drawing of the hair of the head and the pubic region, 
in both of which the locks were slender and had a typical spiral twist. The 
scalp lock closely resembled that of the Andaman islander drawn in fig. 19. The 
individual hairs were oval in transverse section, the longer diameter 105m, the 
shorter 75m, whilst the colour was generally dark brown. 

I now propose to compare the Ulotrichous hair of the Tasmanians, Melanesians 
and Papuans with the Leiotrichous Australians, Maoris and other Polynesians, their 
neighbours in the Pacific region. 

Australians. 

It is generally acknowledged that in the aborigines of Australia the hair is black, 
relatively long, frequently straight, or with coarse natural curls, neither woolly nor 
artificially dressed in pencil-like ringlets, nor expanded into a mop-like mass, nor 
frizzled at the free ends (figs. 21, 22). The University collection is indebted to Dr Wm. 
Ramsay Smith for heads from South Australia on which the hairy scalp had been 
retained. In all, the hair was black and straight, each hair had its independent 
course, and was not artificially dressed. In a man the hair did not exceed 4 cm. in 
length ; in two women the length in one was 6 cm., in the other 10 cm. (4 inches). A 
cutting of hair of a woman from Benalla, Victoria, was in wavy locks, 15 cm. long, 
which could be stretched to 18 cm. (7 inches). Another specimen presented by 
Dr Frederick Page was cut from the scalp of a native of Victoria plains, West 
Australia ; it was a loose curl, 10 cm. long, which could be stretched to 18 cm. 

I am indebted to Mr Ling Roth for cuttings of hair from the aborigines of North 
Queensland, collected by his brother Dr Walter E. Roth. The specimens were 38 
in number, 29 of which were from adult males, 8 from adult females, and 1 from an 
infant three weeks old. They had been obtained from natives of the Morehead, Nor- 
mandy, Jack, M'Torr, Johnstone, Batavian, Musgrave and Lynd River districts, from 

* Tydschrift van het Koninklyk Nederlandsch aardrykskundig Genootschap. 

t "Ueber Pygmaen in Niederlandisch Sud-Neu-Guinea," Zeitschrift fiir Ethnologie, 45th Jahrgang, p. 23, Berlin, 
1913. 

TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 10). 45 



326 PRINCIPAL SIR WILLIAM TURNER ON 

Capes Melville and Grafton, Barrow Point, Princess Charlotte Bay, Palm Isles and from 
the Coen, Herberton, and Chillagoe districts. In all the hair was black, though in some 
deeper in tint than in others. In the men the locks, usually from 4 to 8 cm. long, 
frequently consisted of a single loose curl, though in others the hairs were straight. 
In the women, again, the hair was wavy and loosely curled ; two specimens measured 
about 25 cm. (10 inches) when the curl was extended. The breadth of the curl in 
the specimens varied from 7 to 16 mm. In one specimen the lock, 10 cm. long, was 
formed of straight hair. The hair of the infant, said to be three weeks old, was 4 cm. 
long and showed a tendency to curl. 




Fig. 21.— Australian. 6. 

The specimens of hair from North Queensland shared, therefore, with those which 
I have described from South Australia, Victoria and West Australia in the common 
character of colour, in being either straight, or wavy, or in loose curls. In Dr Walter 
Roth's collection from such stations in Cape York Peninsula as the Normandy, 
Batavian, Musgrave and Lynd districts, the Coen, Capes Melville and Grafton and 
Princess Charlotte Bay, special attention was paid to the characters of the hair, as, 
from the proximity of Cape York to New Guinea, a migration of Papuans to the 
north of Australia might have been possible and a consequent intermixture with the 
Australian aborigines occasioning modifications in the hair. The natives of these 
stations showed, however, no sign of hair frizzly or woolly in its character. 

None of the specimens above referred to gave evidence of artificial dressing of the 
linir. A specimen from a native of Pink River, near Charlotte Waters, presented by 



THE ABORIGINES OF TASMANIA. 



327 



Dr Ramsay Smith, consisted, however, of two locks, 30 cm. and 20 cm. long respec- 
tively. Each was formed of three strands plaited together, and in each strand the 




Fig. 22. — Australian. 



hairs were twisted on each other, the lock was very slender, only 3 mm. broad. 
From the length of the locks the specimen was probably from the head of a woman. 



Polynesians. 

By this term one understands the Maoris of New Zealand and the aborigines of 
groups of islands across the Pacific from Tonga in the west to Easter Island, and 
as far north as the Sandwich Islands. Their chief physical characters are yellowish- 
brown skin, brachycephalic heads, average stature of men 5 feet 9 or 10 inches, hair 
black, long, straight, wavy or curly. 

Maoris. — The University Museum contains six dried heads, the faces of which 
have been tattooed and the hair of the scalp preserved. In one, xxxi. A, No. 51, that 
of a chief elaborately tattooed, the hair was parted along the middle and fell in 
large locks down the sides and back of the head and neck ; at first it was wavy 
and it then formed loose curls which had reached the shoulders (fig. 23). The strands 



328 



PRINCIPAL SIR WILLIAM TURNER ON 



of hair were from 30 to 37 cm. long and constituted a remarkable covering for the 
head. The lips and chin had a well-grown moustache and short beard. Nos. 47, 49 




Fig. 23.— Maori, New Zealand. No. 51. 

and 50 had hair which at first was almost straight but it ended in loose curls. 
In No. 48 the hair was scanty and consisted of a median sagittal strip with a lateral 




Fir;. 24.— Maori, New Zealand. No. 52 



strip on each side following the line of the coronal suture. In No. 52, a female, the 
hair at the parting was thin ; otherwise it was arranged in large loose curls, which, 
when uncoiled, were from 5 to 6 cm. long, but could be elongated to from 10 to 



THE ABORIGINES OF TASMANIA. 329 

12 cm. In another specimen, No. 55, in the Museum, the hair had been removed 
from the scalp. 

Captain Cook described* the Maoris as he saw them in 1770. They were rather 
above the ordinary size, of a very dark-brown colour, the face tattooed, black hair, 
thin black beards and white teeth. In the men the hair was long, sometimes combed 
on to the crown of the head ; t in the women it was at times long, at others cut 
short. Numerous figures of the tattooed heads of Maoris are given by Major-General 
Robley in his profusely illustrated work,| a few from life, though mostly from dried 
heads, in some of which the hair was short and straight ; in others a little longer and 
in large loose curls ; in others long, wavy and ending in loose curls. In no sense 
could the hair be described as short and woolly in the sense in which the term is used 
for the hair in the Negro, or long and frizzled as in Melanesians. 

Easter Island. — This island was visited by Captain Cook in 1774,§ who gave a 
portrait of a man and a woman. He described the hair as black, worn long by the 
women, and sometimes tied on the crown of the head ; the men had the hair of the 
head and beard cut short. They tattooed the skin of both face and body, and in 
colour, features and language bore affinity to the islanders further west. Later 
visitors have also called the hair black or dark brown, straight, smooth or wavy. 
In 1902 Dr A. B. Meyer presented me with a coil of hair from a native of this island. || 
Its colour was dark brown approaching to black ; it was neither curly nor frizzled but 
smooth and silky. Its characters, therefore, were Polynesian and not Melanesian. 
The skin, it is said, was not black, but light brown when not exposed to the sun. 

Hair of Scalp, its Implantation, Form and Structure. 

The hair in man, in its mode of implantation in the scalp and in its form and 
structure, has been well described in text-books on histology. The descriptions have 
been chiefly based on the study of the hair in Europeans. Several copiously illus- 
trated works in which an account is given of the form and structure of the hair in a 
few non-European races, and in certain mammals, have also been published, viz. : — 

W. Waldbykr, Atlas der menschlichen und lierischen Haare, so wie der dhnlichen Fasergebilde, 
Lahr, 1884. 

Hans Friedenthal, " Entwicklung, Bau und Entstehung der Haare," Lieferung IV., Beitrdge 
zur Naturgeschichte des Menschen, Jena, 1908. 

Hans Friedenthal, Tierhaaratlas, Jena, 1911. 

Gustav Fritsgh, Das Haupthaar und seine Bildungsstdtte hei den liassen des Menschen, 
Berlin, 1912. 

* Journal of the First Voyage, edited by Captain Wharton, p. 218, London, 1893. 

t In vol. iii. of Hawkesworte's Edition of Cook's First Voyage (1773), plate xiii. p. 49, a tattooed head is depicted 
with the hair dressed in this way. 

| Moko or Maori Tattooing, London, 1896. Fig. 172 is very like my fig. 23 on p. 328. 

§ Second Voyage, vol. i. Plates Nos. xxv., xlvi., p. 290, 1777. 

|| Abhand. u. Bericht. Kbnig. Zool. u. Anthr. Museums zu Dresden, Bd. ix., 1901. 



330 PRINCIPAL SIR WILLIAM TURNER ON 

Implantation. — An important character in the growth of the hair in the scalp 
was recognised by the late Mr Charles Stewart,* who compared the implanta- 
tion of the hair in the European with that in the Negro, and who stated that 
in the latter the hair and its follicle were " remarkably curved so that they 
commonly described a half circle," whilst in the European the follicular hair was 
straight. This observation on the follicular hair in the Negro was verified by 
Dr Anderson Stuart, t though the amount of the curve was, he said, not more than 
about a quarter circle ; further, he associated the curve with the curl of the hair, 
which gave to the hair in the Negro its woolly character after being extruded 
through the skin. 

Fritsch in his recently published treatise has added materially to our knowledge 
of the implantation of hair and the direction of its follicle in various races of men. 
He has described numerous sections through the scalp, perpendicular as well as 
transverse to the surface. Thus in Europeans, Chinese, Indians, Javanese and 
Fellahs the follicle with its included hair was directed vertically, or almost vertically, 
to the surface of the skin : but in other Europeans, in Fellahs, Arabs, Abyssinians 
and Japs the direction was oblique but not curved. In Admiralty Islanders again 
and other Melanesians, in Papuans, in the Herero Bantu people, and more strongly 
in the Hottentots he saw the follicle with its included hair markedly curved. The 
vertical and in some degree slightly obliquely directed hair follicles were thus associ- 
ated with straight-haired races, whilst the curved follicles were characteristic of the 
woolly and frizzly haired people. The inference drawn by Anderson Stuart from 
his observation on a Negro has therefore been confirmed by more extended enquiries. 

I have examined the implantation of the scalp hair of the Negro in both vertical 
and transverse sections. When the section was perpendicular to the surface the 
hairs and follicles could seldom be seen in their entire length ; they were cut across 
obliquely sometimes more than once. It was obvious, therefore, that the hairs did 
not pass straight from the papilla of origin to the place of exit on the surface, but 
had a curved direction, which varied in its degree in different hairs. It seemed as 
if the curve of the hair increased in rapidity as it approached the surface, so that it 
emerged at a more or less acute angle to the surface of the scalp (fig. 25). 

The deep end of the follicle had a characteristic appearance. Instead of being 
vortical like the papilla it was usually abruptly bent to one side, as if, from the 
beginning, an oblique or curved direction was thus given to the growth of hair and 
follicle, which, together with the more rapid curve near its emergence, contributed 
to the production of the spiral turn of the woolly locks on the scalp of the Negro. 

Fritsch figured the hair in the scalp in several tribes of African Bantus and 
Papuans in which the direction of the hair follicle was like that above described, 
and the bend in the follicle in the Hottentots was so marked that its direction 

* " Note on the Scalp of a Negro," Royal Microscopical Society, January 1, p. 54, 1873, 
t Journal of Anal, and Phys., vol. xvi. p. 362, 1882, 



THE ABORIGINES OF TASMANIA. 331 

near the deep end was almost horizontal, and the scimitar-like curve of the hair 
approached half a circle. Vertical sections through the scalp, which he figured 
in the straight hair of Chinese and Europeans, contrasted, in the vertical or 
slightly oblique direction of the hair follicles, with the curved follicles of the 
woolly-haired races. 

I noted that a sebaceous gland was situated close to each hair follicle about 
opposite the juncture of its middle and upper third, and it reached almost to the 
pigmented Malpighian layer of the epidermis. With a curved follicle the gland was 
next its concave aspect. A band of non-striped muscle, the arrector pili, seen in 
many sections passed obliquely from the superficial part of the cutis to end in the 
hair follicle in proximity to the sebaceous gland. 

The sections through the cutis of the Negro, parallel to the plane of the surface, 
varied in appearance in accordance with the distance from the epidermis. At its 
deepest part the closed ends of the hair follicles were interspersed with coils of sweat 
glands and small collections of fat cells in the connective tissue. The follicles were 
cut across either transversely or obliquely. In some cases the section passed through 
the cells of the root sheath and bulb before cornification, in others the section was 
through the deep end of the hair itself. In each case the root sheath of the hair 
was seen to consist of an outer division lining the follicle, and an inner division 
investing the bulb, or the hair. The bulb, approximately circular in section, was 
composed of small nucleated cells somewhat pigmented. The inner and outer root 
sheaths in the plane of the bulb were of almost equal thickness, but in relation to the 
hair the outer sheath was distinctly thicker than the inner. The cells of the inner 
sheath were nucleated and elongated into short columns. In the outer sheath the 
nucleated cells were more numerous, and those most external formed a distinct layer 
next the wall of the follicle. In this plane of section the follicles with the contained 
hairs were mostly single, though some were in pairs placed side by side (fig. 27). 
Coiled tubes of the sweat glands, with their cellular contents, and fat lobules were 
situated close to the hair follicles. 

In transverse sections, not quite so deep, fewer single hairs were seen and the 
arrangement in pairs was more general. One of the twin hairs with its follicle was 
always distinctly larger than the other. The amount of difference in size may be 
expressed in the following figures : larger twin hair 0'0575, smaller 0'0425 ; larger 
twin 0'06, smaller 0'03. Each hair, or a pair of twins, was enclosed in a ring-like 
circle of connective tissue. Occasionally the section of the hair in its follicle was 
circular, but in most cases the sides were more or less flattened, with one side indeed 
somewhat indented, so that the section was ovoid, or slightly kidney shaped. A 
transverse section of the flattened hairs showed two diameters, one longer than the 
other in accordance with the degree of flattening. 

Sections made immediately below the cuticular surface showed more completely 
the grouping of the hairs. Three hairs in their respective follicles were not uncommon, 



332 PRINCIPAL SIR WILLIAM TURNER ON 

and in rare cases a fourth was included in a group. The third or fourth hair was 
much smaller than the lesser of the two larger hairs, and the papilla from which it 
sprang had not sunk so deeply in the cutis as was the case with the larger hairs. 
In these sections other important changes in the cutis had taken place, the collections 
of fat cells were not present, the coils of sweat glands had disappeared, though the 
excretory ducts could be seen at intervals cut across in their course to the surface, 
and a new and important factor, the sebaceous glands had appeared. These glands 
were grape-like in character and consisted of several vesicles which opened into a 
common duct (fig. 28). The vesicles contained the nucleated secreting cells, some of 
which showed the characteristic granular fatty secretion of the gland. The gland 
duct opened into the funnel-shaped depression which also received the hair as it 
emerged from its follicle. 

The implantation of the hair in the scalp of a straight-haired aboriginal from 
South Australia was then compared with the woolly -haired Negro. In sections through 
the skin, perpendicular to the surface, the hairs and follicles were seen for the most 
part placed vertically and directed straight from the deep part of the cutis to their 
emergence through the epidermis. Some follicles were, however, oblique in direction 
and had consequently been obliquely divided. The follicles were not curved. The 
deep ends of the follicles were not all at the same depth, but as a rule they were in 
direct line with the axis of the hair and the papillae were vertical : in some specimens, 
however, this end was bent a little to one side, so slightly indeed as not to affect the 
curvature of the hair. In many specimens the whole length of the imbedded hair 
was present in its follicle, and at its emergence it was not acutely bent on the scalp 
(fig. 26). If a follicle had been obliquely divided, only a portion of the hair could be 
seen at one time. Most of the hairs in the follicles were marked by dark lines or spots 
as if pigmented, but a small proportion were almost colourless. It was noted that the 
coils of the sweat glands did not lie deeper than the hair follicles, and that in places 
they were found in close relation to the deeper vesicles of the sebaceous glands, the 
latter of which were large and almost reached the Malpighian layer of the epidermis. 

In transverse sections through the deeper part of the cutis in the Australian the 
hair follicles were scattered singly in the connective tissue (fig. 29). Some were cut 
across through the bulb in which the nucleated cells, soft and distinct, had not become 
cornified. In others cornification had taken place. The cells of the two divisions of 
the root sheath were well defined. A few scattered tubes of sweat glands were seen. 
In sections not so deep the follicles and cornified hairs were lying in pairs, one hair 
being somewhat larger than the other ; the sweat glands were more numerous. As 
the sections approached the cuticle the grouping of the follicles became more marked, 
three hairs were usually grouped together, though a much smaller fourth was some- 
times seen. The sebaceous glands were large and close to the follicles, and as the 
intermediate connective tissue was reduced in amount, the section was largely made 
up of the glands and follicles (fig. 30). The gland vesicles were distinct and lined 



THE ABORIGINES OF TASMANIA. 333 

by a well-defined layer of nucleated cells, internal to which numerous fatty particles 
could be seen, contained apparently in cells occupying the interior of the vesicle. 
Sections through the arrectores pilorum were numerous and many were seen attached 
to their respective hair follicles in one, two, or three bundles. In sections immediately 
below the rete Malpighi the sebaceous glands had disappeared, though the ducts were 
present, whilst the hairs were rearranged usually in pairs, prior to their emergence 
through the cuticle. 

Form and Structure. — A hair begins at the bottom of its follicle in a dilatation, 
or bulb, which surmounts and surrounds the papilla. Its follicular length in the scalp 
of the Negro averaged about 2 mm., in that of the Australian 2 '5 mm. ; after its emer- 
gence, as the shaft of the hair, the length varied greatly in different races, but was 
longer in the Leiotrichi than in the Ulotrichi. The shaft diminished in diameter 
towards its free end, and, when uncut, it terminated in a point. 

In the course of time it became known that the shafts of the hairs were not 
uniform in shape ; sometimes they were cylindriform, at others more or less flattened 
at the sides. Fifty years ago Pruner Bey undertook an extensive investigation 
into the form and breadth of the shaft in different races.* He made transverse 
sections of hairs and examined them with the microscope. Similar observations were 
subsequently conducted by M. TopiNARD.t It was thus recognised that some hairs, 
when transversely divided, were circular in form or approximately so ; that others 
again, more or less laterally compressed, were ovoid, elliptical, or kidney shaped. In 
the compressed group the section had two diameters, one longer than the other, and 

a hair index was calculated by the formula — = : . On the basis that 

J 2 

the long diameter was equal to 100, the index of breadth was its relation to that 
number. The hair index was low in the woolly-haired black races, in which the 
section was laterally compressed, and higher in the straight-haired races, in which 
the section approximated more to the circular. Topinard from his own observations 
placed the yellow races in a group the hair index in which ranged from 79 to 90, 
the black races in a group from 40 to 60, whilst in an intermediate group com- 
prising Europeans, Polynesians, Laps, Fins and Australians, the index ranged from 
62 to 74. 

Waldeyer in his treatise figured transverse sections through the hair of the 
head in Europeans, a Negro, a Jap and a Jew : Friedenthal also gave sections of 
hair in Europeans, Chinese, Papuans and in Togo negros. Fritsch in his great 
Atlas represented sections of the hair of Europeans, Esquimaux, the Yellow races, 
Polynesians, Australians, Papuans, Bushmen, Zulus and Negritos. 

Elaborate descriptions of the structure of the hair have been given in the best 
histological text-books4 In the three illustrated special treatises on the hair 

* Mem. Soc. Anthropologic, t. ii. iii., 1863-64. t Elements d'Anthrop. generate, p. 278, Paris, 1885. 

I For example Kolliker, Handbuch der Gewebelehre des Menschen, 1889. 

TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 10). 46 



334 PRINCIPAL SIR WILLIAM TURNER ON 

already referred to (p. 329) careful drawings of the medulla, the cortex and the 
cuticular covering in several races have been reproduced. 

Tasmanians. — Opportunities of examining the form and structure of the hair in 
this now extinct race have seldom occurred. The only references which I have found 
are — (a) statement by Topinard * that the hair was flattened laterally and was ovoid 
or elliptical in transverse section, and that the hair index was 60, which was modified 
by Latteux to 63 ; (b) Sydney Hickson described t the hair as light golden 
brown in colour, curly, very flat in transverse section, the average diameter of the 
curl being 5 mm. ; the Tasmanian hair, he said, was finer than that of the Papuan, 
but not so fine as in the Andamanese ; (c) Friedenthal, in a recently published 
paper,J recorded the dimensions of a small sample of hair which Professor von 
Luschan had supplied to him. The sample consisted of twelve hairs from 15 to 
91 mm. long, without roots but with their natural tips. The hairs, he said, were of 
reddish-brown colour and were curled comparable with the hair of Papuans. They 
were not cylindrical and the breadth in one diameter exceeded the depth in another ; 
a typical breadth was 0'095 mm., a typical depth 0*065 mm. If the breadth or 
greatest transverse diameter is regarded as = 100, the hair index is about 68. 
Comparative measurements of the hair in other races are given by Friedenthal, 
the mean maximum breadth being as follows : Europeans 0'102 mm., American 
Indians 0'09 mm., Chinese 0"099 mm., Japanese 0'105 mm., Bushmen 0'0773 mm., 
and Bantu Hereros 0*083 mm. 

My observations on the form and structure of the hair of the head in Tasmanian 
aborigines were made on samples from two men and the old woman Truganini, 
also on some hairs of Mrs Cochrane Smith of mixed blood (pp. 310-311). In the 
aborigines the hair was black, or brownish black. The hairs obviously tapered to a 
fine point at the tip, which however had been frequently cut. The measurements 
were made with a screw micrometer eye-piece. In one male the root sheath had 
been drawn along with some of the hairs from the follicles ; the breadth of the root 
hair, as well as the shaft was taken ; in this specimen the short hairs referred to on 
p. 311 were also measured. Measurements of the clippings from the other male 
and from Truganini were also taken : — 



Male, maximum breadth of root 


hair 


00875 


mm. 


>> !> 


, shaft 


)> 


0-0700 


)> 


,, clipping of hair 


> . )> 


>) 


00950 


5> 


>) >) 


tip 


j> 


0-0125 


)> 


Truganini 


, shaft 


>j 


0-0575 


)1 


Short hair in male 


, root 


jj 


0-0725 


)) 


i) >) 


, shaft 


>> 


0-0675 


)) 


!) )) 


tip 


>> 


00200 


)) 



* Elements d'Anthrop. ge'ne'rale, pp. 278, 280, 1885. t In Ling Roth's Aborigines of Tasmania, p. 226. 

+ « Vergleich von Tasmanier-kopf haaren mit den Kopfhaaren anderer Menschenrassen," Zeitschr.fiir Ethnologie, 
45. Jahrgang, Heft 1, p. 49, 1913. 



THE ABORIGINES OF TASMANIA. 335 

When the hairs were examined microscopically, the shaft was seen not to be 
uniform in breadth, even in parts in close proximity to each other, owing to their 
resting sometimes on the larger diameter (breadth), at others on the smaller diameter 
(depth), so that it presented different aspects to the observer and was as if twisted 
on itself (fig. 31). The measurements of the two diameters, when computed by the 
formula, gave a hair index which averaged 67* 

I examined the structure of the Tasmanian hair under low and high powers. 
The cortex, or rind, constituted the bulk of the hair, and was marked with dark 
spots and lines running longitudinally, which gave it the fibrous character and black 
or dark brown colour. The margin of the hair was distinguished by a bright, 
narrow, non-pigmented line which represented the thickness of the cuticular covering. 
Occasionally the hair split longitudinally in the direction of its fibro-cells. It was 
exceptional not to see an axial medulla in the dark hairs, both in the shaft and the 
follicular end ; and the medulla constituted about one-fifth of the breadth of the hair. 
In many cases the medulla formed a continuous band for a large part of the length 
of the hair, but in others it was broken into short divisions or fragments. The 
medulla was so dark that its constituent cells could not be recognised. I have 
called attention on p. 311 to short delicate hairs emerging from the scalp. When 
closely examined they were seen to be brown, with a clear cuticle and dark lines 
and spots in the cortex, the latter of which were sometimes diffused ; no sign of an 
axial band of medulla was visible. 

I have referred on p. 311 to the hair of Mrs Cochrane Smith, a Tasmanian half- 
breed, which differed both in colour and character from a pure aboriginal. Under the 
microscope two kinds of hair were distinguished, the one in shades of brown, mostly 
pale in tint, the other practically colourless. In neither was a pigmented band of 
medulla seen, and the hair consisted of cortex with a cuticular covering. The brown 
colour was clue to spots and longitudinal lines in the cortex. The hair shaft varied 
in breadth, though without a definite twist, the maximum being about 0'08 mm. 
and the minimum about "050, which gave an index about 65. 

New Hebrides. — The observations were made on hair cut from the scalp. The 
shaft examined microscopically by transmitted light was not uniform in character, a 
broader diameter alternating with a narrower, owing to the twist of the hair on 
itself. In the middle of the shaft the broader part ranged from "097 to '08 mm., 
and the narrower part from "0625 to "0525, which gave a mean hair index 73. The 
cut ends of the hairs were from "055 to '08 mm. ; the diameter of the tip of a 
hair was '025. In structure the cortex had the customary fibrous appearance in 

* It has been customary to compute the hair index from measurements of the larger and smaller diameters in 
transverse sections of the hair. In determining the index in the Ulotrichi I have measured the larger and smaller 
diameters in adjoining parts of the twisted hair shaft itself, without making sections, and have computed the index 
with the customary formula. The results closely corresponded with those obtained by other observers from the 
measurement of transverse sections. The index of the hair in its follicle was, however, computed from transverse 
sections of the hairs as they lie in the cutis. 



33() PEINClPAL SIR WILLIAM TURNLH ON 

shades of brown. In each hair examined a distinct black axial medulla was present, 
which frequently extended for a considerable distance unbroken, but sometimes it 
was interrupted by relatively long intervals, though at others with only short 
gaps, and the cortex constituted the body of the shaft. In places the medulla 
was so broken up that the outlines of the cells could be seen. 

Solomon Islands. — The hairs were mostly coated with a black tenacious material, 
which was removed with difficulty, and until this was taken off the hair was black 
and opaque. The cortex of the hair was then seen to be dark brown with pigmented 
spots and longitudinal dark lines. In some hairs only cortex and cuticle were distinct, 
but others possessed a black medulla which was frequently broken into short sections. 
The diameter of the shaft ranged from '0500 to '0750, and that of the tip was '0200. 

New Guinea. — The specimens were cuttings. In the locks from Rubi, Geelvink 
Bay, the hairs were twisted; the broader diameter of the shaft was about '07 mm., 
the narrower '05 mm., the hair index about 63 (fig. 10). The cut ends measured 
in breadth about '05 ; the diameter of the pointed tip of one hair measured was 
'02 mm. The individual hairs gave characteristic views of the light to dark brown 
fibrous cortex, and a dark axial medulla was also present. The medulla had a 
strong disposition to break up into fragments, so small indeed in places that the 
individual cells could be recognised without difficulty. The cortex was occasionally 
split longitudinally. 

In the cuttings of human hair from a dance mask from the Papuan Gulf the cut 
ends were on the average about '05 mm. in breadth, but the free tip of a hair was 
only '0225 mm. The shaft showed the customary broader and narrower alternations 
in diameter due to the twist present in woolly-haired races ; measurements ranging 
from '0875 to '0525 were noted, and a mean hair index 70 was computed. Under 
the microscope the hairs varied in their depth of colour, some were so opaque that 
their structure was ill defined, others were a deep brown, others a much lighter 
brown. In those lighter hairs the fibrous structure of the cortex was recognised, 
in some of which an axial dark medulla somewhat interrupted was distinct, whilst 
others again showed no trace of medulla. 

Negro. — In transverse sections through the cutis of the scalp, I measured the 
diameter of several hair follicles which ranged from 0'1 mm. to '0275, so that in 
the same plane of section a great diversity in the breadth of the follicle was seen. 
The divided cornified hairs lying in the follicles were also measured. It was excep- 
tional to find one circular in outline or even approximately so. As a rule they 
had two diameters, one definitely longer than the other, so that the hair was 
flattened laterally and the section was either a wide or an elongated ovoid, with 
sometimes a kidney-like depression on one aspect. The breadth or longer diameter 
of the section of hair in the larger follicles ranged from '075 to '0375, whilst the 
depth or shorter diameter was from '045 to '0325 mm.; the index computed from 
these dimensions was 51. The ends of the shafts cut from the scalp averaged 



THE ABORIGINES OF TASMANIA. 337 

'056 in breadth and the broadest was '07. Each hair, when lying on a microscopic 
slide, had inequalities in breadth, even in closely adjoining parts of the shaft, 
owing to the natural twist in a hair, and the index computed from the longer and 
shorter diameters was 53. 

As a rule the hairs were so intense a black that the structural characters were 
obscured. Sometimes a brownish-black shade permitted transmission of a little 
light reflected from the mirror. The fibrous structure of the cortex was expressed 
by dark lines and spots, and by an occasional longitudinal splitting of the hair 
in the course of the fibres. An axial band of dark medulla was not recognised 
in any specimen, but the margin of the hair was defined by the cuticular 
covering. 

Bush. — In the hairs of this race the alternating broader and narrower diameters 
of the shaft were well pronounced, and the broader diameter ranged from '0625 to 
'05 min., whilst the narrower was from '05 to '035 ; the hair index was computed to 
average 50 (fig. 32). The cut ends of the hairs measured from '035 to '05 mm. The 
colour of the hair was black, though modified at times with a brown tint ; it pervaded 
the whole cortex with coarse lines and spots, but I saw no definite axial medulla. 
The grey hair of the old woman presented interesting features, for whilst the 
majority of hairs were colourless and the lines in the cortex were very feeble, 
a few had a distinct axial medulla. In some hairs the medulla, though almost 
colourless, was distinct from the cortex and showed indications of cells, but in others 
it was black and strongly differentiated. A few normally coloured non-medullated 
hairs were interspersed amongst the grey hairs. 

Hottentot. — The broader diameter of the twisted hair shaft in the specimens 
ranged from '0625 to '05 and averaged '0580 mm., whilst the narrower diameter 
ranged from '045 to '03 and the mean was" '038 mm. ; the hair index was com- 
puted at 50. The cut ends varied in diameter from '0525 to '0375 mm. The bulk 
of the hairs were black and so opaque that the structure was obscure, but a proportion 
had different shades of brown. The cortex showed its customary characters and 
was sometimes split longitudinally. A black axial medulla was frequently seen 
either as a continuous band, or in fragments, some of which were so small that they 
might represent individual pigmented cells. 

Kaffir. — The shafts of the twisted hairs showed a broader diameter which had the 
mean '0616, and a narrower '0437 mm. ; the computed index was 52. The cut ends 
ranged from '04 to '0350, and the diameter of the tip was '0225 mm. The cortex 
was so deep a black that the structure was obscured, but some hairs were split 
longitudinally in the direction of their fibres : an occasional brown-black hair was 
seen ; it is difficult to say if a medullary band was present, though probably not. 

Andaman Islanders. — The cuttings of hair of these Negritos, as they rested on 
microscopic slides, showed a marked twist, and the passage in the shaft from a 
broader transverse diameter to an adjoining narrower could be easily traced. The 



338 PRINCIPAL SIR WILLIAM TURNER ON THE ABORIGINES OF TASMANIA. 

broader ranged from '085 to "0625 with a mean '0725 mm., whilst the narrower 
was from '0525 to '0475 with a mean '050 mm. The hair index was 61. The 
cut ends ranged in breadth from '08 to "035. In a few hairs the shaft began with the 
hair bulb, the largest example of which was "130 in transverse diameter : in another 
specimen the tip of the hair was '0225 in breadth. 

When examined by transmitted light the hairs showed various shades of brown, 
but were not black. The cortex was characteristically marked with brown spots 
and lines, and had a cuticular covering. The medulla was not recognisable as a 
black band, though in some hairs an axial stripe could be seen not deeper in tint than 
the cortex itself. 

Semang. — The specimen consisted of a single short lock curled so as to form a 
spiral. The shafts of the individual hairs were twisted and showed variations in 
breadth in adjoining parts of the length ; the broader diameters ranged from '075 to 
'0575, with a mean '0630 mm., the narrower '0375 to '0350 with a mean "0362 mm. ; 
the hair index computed was about 50. The diameter of the cut ends ranged from 
'07 to '0350. The hairs as a rule were black, but with transmitted light a few brown 
hairs were seen interspersed in the lock. The opacity of the black hairs interfered 
with the determination of structure. In the brown hairs the cortex was characteristic 
and an occasional hair was split in the direction of the fibres. In some brown hairs an 
interrupted axial medulla was present in which dark-brown cells were recognised. 

Australians. — Transverse sections through the hairy scalp of the South Australian 
enabled the form of the hair in the follicles to be seen. Some were cylindriform, the 
majority had in section two unequal diameters with an ovoid outline. The longer 
diameter ranged from '0925 to '0675 mm. in breadth, the shorter from '06 to '0475 mm. 
in depth. One of the largest hair bulbs across the papilla was '112 mm. and the 
broad diameter of one of the smaller follicular hairs was only '0350 mm. Hairs 
measured at their cut ends and in the shaft showed variations in breadth and ranged 
from '107 to '0475, the mean of the longer diameter was '0860, that of the shorter 
'0595, and the computed index of the shaft was 72. The narrow end of a hair near 
the tip was '0425. 

The shaft of an individual hair was smooth and almost uniform in transverse 
diameter (fig. 33, A) until it began to taper towards the tip. As a rule the hairs were 
black and opaque, though a few were deep brown and permitted a transmission of light. 
The cortex and its cuticular covering were usually the sole constituents of the brown 
hairs ; but, in a small proportion, a medulla, darker in colour than the cortex, occupied 
the axis, and could be traced as a continuous band for some distance, though in places 
it broke into shorter fragments. 

Maori. — The hair of the New Zealand aborigines was smooth, not twisted, and was 
almost uniform in breadth in the greater part of the shaft, amounting frequently to 
*l mm. or a little less (fig. 33, M) ; the mean of the cut ends through the shaft was '08, 
and the lowest transverse diameter was '065 mm. ; the hair index was computed as 85. 





''ig. 25. — Negro. V.S. 



Fig. 26.— Australian. V.S. 







Fig. 28.— Negro. T.S 



Fig. 30.— Australian. T.S. 



340 PRINCIPAL SIR WILLIAM TURNER ON 

The hairs were usually black and so opaque that the structure could not be accurately 
differentiated ; sometimes, however, hairs permitted some light to be transmitted, 
when either no medulla was seen, or occasionally a black medullary band, enclosed 
by the less opaque cortex, formed the axis of the hair. 

Easter Island. — The shaft in the individual hair was uniform in breadth, until it 
began to taper to the tip ; it showed no trace of a twist such as one sees in the Ulotrichi, 
and was a good example of a straight smooth-haired race. The breadth of the shafts 
and of their cut ends averaged about '07, the shaft in its broadest part ranged from 
"07 to '087 and it was not twisted. The hairs were in shades of brown, though in 
mass they were black brown, and the dark spots and lines in the fibrous cortex were 
well marked. Many hairs consisted of cortex with its cuticular covering, but in others 
a dark axial medulla could also be seen, which had a tendency to break up into small 
divisions (fig. 33, E). 

Summary. 

1. The locks of hair in the Ulotrichi are characterised by being either short and 
woolly, or moderately long and frizzly. Three factors combine to produce hair of 
this character : 

(a) A curved, and not a straight, follicle in the cutis ; and a consequent curve 
of the contained hair, which emerges at an acute angle to the surface of the scalp. 

(b) The hair shaft in transverse section is not cylindrical, but oval, elliptical, or 
kidney-shaped, one diameter of which forms the maximum breadth of the shaft, whilst 
the shorter diameter at the end of the section expresses the depth of the section. 

(c) Each hair shaft is twisted on itself, and when placed on a flat surface it lies 
alternately on its lateral maximum diameter and on its shorter diameter. 

(d) The individual hairs after their emergence collect into locks in which the 
hairs are spirally curled. When the spiral is strongly compressed the locks arc 
short and woolly ; when it is more open the locks are longer and more like narrow 
ribbons or cords. 

(e) The hair is either black or brownish black, though sometimes it is artificially 
bleached to a light or yellowish-brown tint. 

2. The Tasmanians in the character of the hair belonged to the Ulotrichi. The 
hair shafts flattened laterally had broader and shorter diameters, the difference 
between which is expressed by a moderate hair index ranging from 63 to 68. The 
shaft was not a narrow band continued in one plane, but was twisted on itself, was 
oval or elliptical in transverse section, and rested, when lying on a plane surface, 
alternately on its broader and shorter diameters. The hairs, as attached to the seal]), 
wiie at times moderately long and arranged in slender ringlet or cord-like locks, in 
which they formed a compact spiral coil, though, when the hair had been cut short, it 
presented the appearance of matted curls ; in colour it was black or brownish black. 
Ajs in only one of my specimens a portion of the scalp had been preserved, and that 



THE ABORIGINES OF TASMANIA. 



341 



was dried and with the cutis imperfect, the form and direction of the follicle with its 
contained hair could not be precisely denned, but from the character of the hair itself 
there can, I think, be no doubt that the follicles and the contained hair were curved, 
as is the case in the Negro, and, as has been shown by Fritsch, in other Ulotrichi — 
Melanesians, Papuans, Hottentots, and Bantus. 

3. The cortex formed the essential constituent of the Tasmanian hair shaft. 
Sometimes no medulla was present ; but, as a rule, the axis of the hair consisted of a 
band of dark-coloured medulla, which might be either continuous or in fragments. 




Fig. 31. — Tasmanian. 




Fig. 32.— Bush. 



The twist in the individual hairs occurred more frequently in a given length of the 
shaft in a woolly hair, as in the Bush, than in the same length of the frizzly-haired 
Tasmanian (figs. 31, 32). 

4. The hair index in the Tasmanians, 63 to 68, was higher than is found generally 
in those Ulotrichi who are markedly woolly. In the specimens measured in this 
memoir the following average indices were obtained : — Negro, 53 ; Bush and Hottentot, 
each 50 ; Kaffir, 52. Of the Negritos the index in the Semang was 50 ; in the 
Andaman Islander, 61. 

5. The Melanesian natives of New Guinea and the New Hebrides had a higher 
hair index, which reached 70 in the Papuans and 73 in the New Hebrideans. They 
approximated in this respect more closely to the Tasmanians than to the Negroes. 
So far as the maximum breadth of the hair shaft affected its texture and gave to it 
a relatively coarse character, the Papuans, New Hebrideans, and Tasmanians reached 
'09 and '08 mm., whilst in the proper negroid races '07 mm. was the maximum 
measured. 

6. It has been generally recognised that in the Australian aborigines the hair 

had the distinctive character of the Leiotrichi ; either straight or wavy, and 
TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 10). 47 



342 



principal sir William turner on 



relatively long ; in which respects it differed from the Ulotrichi. But in other 
characters it also showed marked differences. In its mode of implantation in the 
scalp the follicle was not curved as in the latter, and with its included hair was 
directed vertically in the cutis. At its emergence it did not make an acute angle 
with the scalp. The shafts of the hairs retained their direct and independent courses 
and did not show a disposition to aggregate into locks, in which the hairs were 
spirally arranged, though they might become wavy and form loose curls. The 
form of the shaft, which was a continuous smooth plane and was prolonged to the 
tip without being twisted on itself, presented also a striking contrast to the woolly 
and frizzly haired races. In these particulars, therefore, the hair of the Aus- 
tralians, as also that of the Maoris and other Polynesians, did not possess those 
developmental and morphological characters which contributed to give to the hair 
of the Ulotrichi distinctive features (fig. 33). 




Fig. 33. — A, Australian ; M, Maori ; E, Easter Islander. 



7. The hair of the Maoris and of the natives of Easter Island, representatives of 
the Polynesians, was relatively long, wavy and sometimes in loose curls, but not 
twisted on itself, or arranged in spiral locks. I have had no opportunity of ex- 
amining the implantation in the cutis, but it may reasonably be inferred that the 
follicles and follicular hair were not curved, but were straight and vertical, or 
approximately so, to the surface of the scalp as in other Leiotrichi. 

8. In Parts I. and II. of the series of Memoirs on the Tasmanians their crania 
were compared with those of Australians, Papuans, Melanesians and the Negritos 
of the Andaman Islands, and the much vexed question of the descent of the 
Tasmanians was considered. 

!). The opinion of anthropologists generally that the aboriginal Australians, as 
we now know them, have marked racial characters, which distinguish them from 
the Tasmanians, is strengthened by the study of the hair described in this memoir. 

10. Differences in the characters of the crania of the Tasmanians, when compared 
with the Papuans and Melanesians, led me to say that the Tasmanians were not in 



THE ABORIGINES OF TASMANIA. 343 

direct descent from the Melanesians as we know them at the present day, though in 
the structure of the hair, as now described, they have some characters in common. 

11. From the comparison with the Negritos of the Andaman Islands I stated that, in 
physical characters, the Tasmanians were more closely allied to the Negritos than to the 
Melanesians. To these I can now add the form and structure of the hair, which, when 
permitted to grow, had in both races a definite arrangement in spirally curled locks 
of moderate length, which might assume the appearance of slender cords or ringlets. 

12. The contrast between the Andaman Islander and the Papuan or Melanesian 
is very marked. In the former the skull was small, rounded, brachycephalic ; parietal 
eminences prominent ; vault flattened ; cranial height less than breadth ; forehead 
smooth, not retreating ; glabella and superciliary ridges feeble : nasion shallow ; 
nasal index platyrhine ; facial profile relatively vertical ; stature dwarf-like ; skin 
dark, approaching black ; hair in short, woolly, slender cord-like ringlets. On the 
other hand, in the Papuan and Melanesian, the skull was long, relatively narrow, 
markedly dolichocephalic ; vault roof-shaped ; cranial height more than the breadth ; 
forehead retreating ; glabella and superciliary ridges strong ; nasion deep ; nose 
platyrhine ; facial profile prognathic ; stature moderate ; skin blackish, or dark 
brown ; hair, when allowed to grow, mop-like and frizzly. 

13. The Tasmanian in his physical characters seemed to occupy a place intermediate 
to the Negrito and the Papuo-Melanesian, though more closely associated with the 
former. The mean cephalic index was sub-dolichocephalic, on the confines of the 
dolicho- and mesaticephalic groups ; the parietal eminences were prominent ; the vault 
had special characters of its own,* though flattened as in the platychamsecephalic 
Negrito ; the hair also closely corresponded to the Negrito. On the other hand, the 
forehead, glabella, superciliary ridges were more in accordance with the Melanesian, 
and the Tasmanian was not a pygmy, but was moderate in stature. 

14. In discussing the question of the origin and descent of the Tasmanians, it 
would be wrong to regard them as a race which had no affinities with any other race. 
From the fact that Tasmania had originally been continuous with Australia, its 
population had undoubtedly been derived from a race coming from the north, 
east, or west and not from the south. Flower regarded the Tasmanians known 
to us, as aberrant members of the Melanesian group showing variations in their 
physical characters, which might have arisen and become intensified and perpetuated 
from their geographical position and insulation. Huxley considered them to be 
a Negrito modification of the Negroid division of mankind, which had migrated from 
Asia eastwards as far as New Caledonia and subsequently southwards to Tasmania. 
Ling Roth spoke of them as aborigines of Australia who had in course of time been 
displaced by the existing straight-haired Australian natives. 

15. I stated in Part I. that the evidence favoured their descent from a primitive 
Negrito stock, which had migrated across Australia, rather than by the route of the 

* Described in detail in Parts I. and II, 



344 PRINCIPAL SIR WILLIAM TURNER ON 

Oceanic islands lying to the north and east of the Australian continent. Since it 
was published an important link in the chain of evidence has been supplied in the dis- 
covery by the English and Dutch explorers in the mountains of West New Guinea 
of a pygmy race possessing undoubted Negrito characters (p. 324). It has therefore 
been established that Negritos continue to occupy limited tracts in the mountains 
of that island, though the country generally is now mostly populated by the more 
virile Papuans. The descriptions by Captain Rawling, Mr Wollaston, and Messrs 
de Kock and v. d. Broek # have satisfactorily established the physical differences 
between the Pygmies and the Papuans, and it is clear that up to the present day a 
characteristic Negrito race has been preserved in the great island to the immediate 
north of Australia. 

16. The possibility, therefore, of the extension of Negritos throughout Australia, 
before its southern end had become an island separated by the Straits of Bass, has been 
accentuated by this fact. The fauna of Tasmania, as regards many species, is identical 
with that of Australia. The human element, when Tasmania was discovered, was, 
however, not the same in the two countries. On the hypothesis that Australia had 
originally been peopled by Negritos, that race had disappeared from it, and the 
straight-haired race, now known to us as the aborigines, had taken its place. The 
fact that similar straight-haired aborigines had not been found in Tasmania, points to 
its separation as preceding their extension into the present southern seaboard of the 
great Australian continent. 

17. The structural differences between the Negritos and the Tasmanians may 
have been due to the geographical insulation of the latter, the production of varia- 
tions from time to time, and their intensification and perpetuation through many 
generations. 

18. The hypothesis that the Negritos had migrated eastwards along the chain of 
islands in the Western Pacific as far as New Caledonia, and had become the ancestors 
of the Melanesians, does not harmonise with the existence, at the present day, side 
by side in New Guinea of Negritos and Papuans. The food, climate and altitude 
in the districts which they respectively occupy are essentially the same, and no 
possible difference in environment would adequately account for the production 
of the modifications in physical structure and in character, which distinguish the 
energetic, sea-loving Melanesian from the shy, forest-retiring, pygmy Negrito, such 
as would lead one to think that they were descended from a common ancestor. 

11). In regard to the ancestry of the Melanesians some anthropologists maintain 
that they have ethnic relations with the Indonesians, who occupy the great islands 
of the Archipelago south of the continent of Asia. It is, however, to be kept in 
mind that in the Indonesians the skull is either dolicho- or mesaticephalic ; the breadth 
and height of the cranium are almost equal ; the mean nasal index ismesorhine ; the 
facial profile is orthognathous ; the stature is low though not that of a pygmy ; the 

* Referred to on p. 325 and in Bibliography. 



THE ABORIGINES OF TASMANIA. 



345 



skin is brown rather than black ; the hair is black, long and straight. Collectively 
these characters constitute marked physical differences from the Melanesians, and 
approximate the Indonesian more to the Polynesian race of the Pacific. 

20. The difference between the direction of the follicle and hair in the scalp, 
and in the form of the shaft, seen in the straight-haired Australian and the woolly- 
haired Negro, applies also without doubt to other races which possess one or other 
of these types. It adds, therefore, an important character in the differentiation of 
Australians from Tasmanians, and in its general application also emphasises the value 
of the study of the development of the hair and the form of the hair shaft, in con- 
sidering the affinities and descent of the races of men. It also raises the question if 
a race belonging to one of these two marked types can be descended from a race in 
which the other type of hair is a distinctive feature. If this were confirmed, it would 
tend to negative the suggestion recently advanced by Sergi, that the descent of the 
Tasmanians has been (through the acquisition of new characters and by the loss of 
others during thousands of years) from a branch of his so-called American genus 
Hesperanthropus, which he thought became diffused over the Pacific area, where, in 
addition to the frizzly- haired Melanesians and Tasmanians, the straight - haired 
Polynesians constitute also an important ethnic element. 




Fig. 34. 



346 PRINCIPAL SIR WILLIAM TURNER ON 

BIBLIOGRAPHY. 

In Part I. of my series of Memoirs on the Tasmanians, published in the Transactions of this Society in 1908, 
I compiled a bibliography of these people up to that date. Since then additional memoirs have 
appeared, the titles of which are subjoined. 

Klaatsch, Hermann, "The Skull of the Australian Aboriginal," Reports of the Pathological Laboratory of 
the Lunacy Department of N.S. Wales, vol. i. pt. iii., Sydney, 1908, in which the skull of a 
Tasmanian in the Sydney Museum is described. 

Berry, R. J. A., and A. W. D. Robertson, "Preliminary Communication on Fifty-three Tasmanian Crania, 
of which forty-two are described for the first time," Proc. Boy. Soc. Victoria, xxii., New Series, 
pt. i., 1909. Anatomischer Anzeiger, Bd. xxxv. No. 1, p. 11, Aug. 10, 1909. "Place in Nature of 
the Tasmanian Aboriginal as deduced from a study of his Calvarium," Part I., Proc. Roy. Soc. 
Edinburgh, xxxi., 1910; Part II., xxxiv. p. 144, 1914. "Dioptrographic Tracings in four Normse 
of Fifty-two Tasmanian Crania," Trans. Roy. Soc. of Victoria, vol. v. pt. i., Dec. 1909. Along 
with K. Stuart Cross, "A Biometrical Study of the Relative Degree of Purity of Race of the 
Tasmanian, Australian and Papuan," in Proc. Roy. Soc. Edinburgh, xxxi. p. 17, 1910. 

Turner, William, " The Aborigines of Tasmania : Part II. The Skeleton," Trans. Roy. Soc Edinburgh, 
xlvii., 1910 : also Part III., " The Hair of the Head compared with that of other Ulotrichi, and with 
Australians and Polynesians," idem, vol. 1., 1914: also, "A Contribution to the Craniology of the 
Natives of Borneo, the Malays, the Natives of Formosa and the Tibetans," idem, xlv. 

Basedow, H., "Der Tasmanier-schadel, ein Insulartypus," Zeitscli. fur Ethnologie, xlii., 1910. 

Biasutti, R., "I Tasmaniani come forma di isolamento geografico," Archivio per V Antropologia, vol. xl., 1910. 

Luschan, Felix v., "Stellung der Tasmanier im Anthropologischensystem," Zeitsch.f. Ethnol., 43. Jahrg., 
1911 ; also, " Tasmanier-Haarprobe," idem, p. 271, 43. Jahrg., 1911 ; " Noch einmal zur Stellung 
der Tasmanier im Anthropologischensystem," idem, p. 123, Heft 1, 1912. 

Merkel, Friedrich, "Tasmanier und Australier," Zeitsch.f. Ethnol., p. 120, Heft 1, 44. Jahrg., 1912. 

Fritsch, Gustav, "Verwertung von Rassenmerkmalen fair allgemeine Verglek-hungen," Zeitsch.f. Ethnol., 
43. Jahrg., p. 272, 1911. 

Poutrin, Or, " Les Negrilles du Centre Africain [Type Brachyc^phale]," U Anthropologic, p. 435, xxi., 1910 
[Type Sous-Dolichocephale], idem, xxii., xxiii., 1911-12. 

Sergi, G., "Tasmanier und Australier — Hesperanthropus tasmanianus," Archiv fur Anthropologie, N.F., 
Band xi. Heft 3, p. 201, 1912. 

Buchner, L. W. G, " An Investigation of Fifty-two Tasmanian Crania by Klaatsch's Craniotrigonometrical 
Methods," Proc. Roy. Soc. Victoria, xxv. p. 122, 1912 ; also, " A Study of the Prognathism of the 
Tasmanian Aboriginal," idem, p. 135 ; also of the Curvatures, P.R.S. Edinb., xxxiv. p. 128, 1914 ; 
also, "Notes on Certain of the Cape Barren Islanders, Bass Strait," Zeit.f. Ethn., 45. Jah., 1913. 

Friedenthal, Hans, " Ueber die Behaarung der Menschenrassen und Menschenaffen, Anhang," Zeitsch. f. 
Ethnologie, 43. Jahrg., p. 979, 1911 : also, "Vergleich von Tasmanier-kopfhaaren mit den Kopf- 
haaren anderer Menschenrassen," idem, 1913, p. 49. 

Neuhauss, Rich., "Ueber die Pygmaen in Deutsch-Neu-Guinea und ueber das Haar der Papua," Zeitsch. f. 
Elhnnlogie, p. 280, 43. Jahrg., 1911 ; also, "Bemerk. zu dem Aufsatz von van dem Broek ueber 
Pygmaen," idem, p. 45, 45. Jahrg., 1913; also, "Das Rothblonde Haar der Papua," idem, p. 259. 

Wollaston, A. F. R., Pygmies and Papuans: the Stone Age and To-day in Dutch New Guinea, London, 1912. 

Rawling, C. G., Land of the New Guinea Pygmies, London, 1913. 

Broek, van dem, A. J. P., " Ueber Pygmaen in Niederlandisch Sud-Neu-Guinea," Zeitsch. f. Ethnologie, 
45. Jahrg., p. 23, 1913. 

Virciiow, Hans, "Buschmannkopf " (head preserved in formalin, showing locks of hair), Zeitsch.f. Ethnologie, 
p. 644, 45. Jahrg., 1913. 

Finsch, Otto, " Ueber Bekleidung, Schmuck und Tatowirung der Papuas der Sudostkiiste von Neu Guinea" 
(modes of decorating the hair are described), Mitlheil. der Anthropo. Gesellsch., in Wien, xv. B., 1885. 

Jens, W. L., " De Papoea's der Geelvinksbaai," Handeling. van de Nederldndsch. Anthropo. Vereenigivg, 
No. 2, June, 1904. 

Noetling, Fritz, " Beitraige zur Kenntnis der archaeolitischen Kultur der Tasmanier," Zeitsch. f. Ethnol., 
43. Jahrg., 1911. 



THE ABORIGINES OF TASMANIA. 347 

EXPLANATION OF FIGURES. 

The locks are specimens of the hair drawn by Mr J. T. Murray: the micro-photographs are by 
Mr Ernest J. Henderson of the Anatomical Museum, from sections made by him; the 
process blocks are by D. Stevenson & Co. 

Fig. 1. Reproduction of portrait of Truganini, the last Tasmanian woman. From a plioto. presented to 
me in 1888 by Dr Lloyd H. Oldmeadow. 

Fig. 2. Small lock of hair of Truganini ; also lock of hair from a male Tasmanian attached to a fragment 
of the scalp. 

Fig. 3. From a photograph. Bust of a male Tasmanian presented to the Museum by Professor 
Anderson Stuart. 

Fig. 4. Natives of Tanna, New Hebrides. From a photo, presented by the Rev. J. H. Lawrie. 

Fig. 5. Three spiral locks of hair from a native of New Hebrides. 

Fig. 6. Locks from a native of Tanna, with some matted hair. 

Fig. 7. Locks from a native of Futuna, New Hebrides. 

Fig. 8. Bunch of hair arranged as a top-knot, probably from a man of the New Hebrides. 

Fig. 9. Native of Port Moresby, New Guinea, showing mop of hair. From a photo. 

Fig. 10. Locks of hair from native of Rubi, Geelvink Bay, New Guinea. 

Fig. 11. Lock from a dance mask, Papuan Gulf. 

Fig. 12. Norma verticalis, head of Negro, showing mat of hair. Photographed from nature. 

Fig. 13. Locks of spiral hair from two Negros. 

Fig. 14. Bushwoman, showing the woolly hair on scalp. Photo, presented by Dr J. B. Lester. 

Fig. 15. Matted hair and separate spiral locks of Bush race. 

Fig. 16. Plaited hair of Hottentot. 

Fig. 17. Zulu Kaffir woman, showing woolly hair on scalp. From a photo. 

Fig. 18. Spiral locks of hair of Kaffir woman. 

Fig. 19. Spiral locks of hair of Andaman Islander. 

Fig. 20. Spiral locks of hair of Semang, from Malay Peninsula. 

Fig. 21. A male Australian with the hair in loose curls. 

Fig. 22. A female Australian with the hair straight or wavy. Figs. 21 and 22 are from photos, pre- 
sented by Dr Wm. Ramsay Smith. 

Fig. 23. Head of Maori chief. From specimen No. 51 in the Anatomical Museum. 

Fig. 24. Head of female Maori. From No. 52 in the same Museum. 

Figs. 25-33. From micro-photographs. 25. Vertical section through scalp of Negro, showing a curved 
follicle with its included hair, the papilla and the hair bulb : the duct of the sebaceous gland opened into 
the funnel-shaped mouth of the follicle. 26. Vertical section through scalp of an aboriginal Australian, with 
straight follicles and hairs, also bulbs, papillae and sebaceous glands. 27. Transverse section through deep 
part of cutis of Negro, showing follicles with contained hairs and root sheaths. 28. Transverse section 
through cutis of same, nearer the surface, showing grouping of follicles, hairs with sebaceous glands ; the 
blank areas are the empty vesicles of the sebaceous glands. 29. Transverse section through deep part of 
cutis of Australian, showing follicles, root sheaths, hairs and sweat glands. 30. Transverse section through 
superficial part of cutis of same, showing grouping of follicles, hairs with sebaceous glands and the arrectores 
pilorum ; the connective tissue is relatively small in amount ; a sweat gland lies at the right border, and 
the sebaceous glands are alongside the follicles. 31. Hairs of male Tasmanian, showing the twist of the 
shaft. 32. Portion of lock of Bush hair, showing alternation of the broader and shorter diameters, due to 
twisting of shaft. 33. A, single straight hair of Australian, M, of Maori, E, of Easter Islander, not twisted. 
A, and M, x 50 ; E. x 90. 

Fig. 34. Group of Andaman Islanders along with Dr Joseph Dougall. From a photo. 



349 ) 



XL— The Pinna-Trace in the Ferns. By R. C. Davie, M.A., B.Sc, late Robert 
Donaldson Research Scholar in the University of Glasgow, Lecturer in Botany 
in the University of Edinburgh. Communicated by Professor I. Bayley 
Balfour, F.R.S. 

(MS. received May 4, 1914. Head June 1, 1914. Issued separately August 18, 1914.) 

(Plates XXXIII.-XXXV.) 

The relation of the leaf-trace to the vascular system of the stem in the Ferns has 
been exhaustively investigated during the past fifteen years by various workers, 
especially by Gwynne-Vaughan and Boodle in this country and by Jeffrey and his 
pupils in America. The interpretations of the numerous types of Fern stele and 
leaf-trace have caused much discussion. And the discovery of a wonderful series of 
fossil Ferns has not settled the discussion of the evolution of the Filicinean vascular 
system, but has carried the battle on to another field. 

The fossils have recently yielded up the structure of their pinna-traces (Kidston, 
'08 ; Bertrand, '09 ; Gordon, '11). Simultaneously an attack has been made on the 
pinna-traces of living Ferns. Various pinna-traces were described in curious mathe- 
matical formulae, but no comparisons were made of the different types, by Bertrand 
and Cornaille ('02) in their Etude sur quelques caracteristiques de la structure 
des Filicinees actuelles. Tansley in 1908 drew attention to the Fern leaf as the 
stronghold of many anatomical problems, and reproduced in detail almost the only 
account of the departure of the pinna-trace, that of Matonia pectinata, R. Br., by 
Seward ('99). Compton in 1909 described the branching of the leaf of Matonia 
sarmentosa, Baker, and figured the departure of the pinna-traces. Chrysler in 
1910, in making comparisons among the Ophioglossacese, referred to pinna-traces in 
the Osmundacese and Polypodiacese, while Sinnott in the same year described the 
pinna-traces of some of the Osmundacese. In 1911 Sinnott made a rapid survey of 
the Filicinean leaf-traces, briefly referring to the relation between them and the 
pinna-traces. Gwynne-Vaughan ('11) amplified some of the American work on the 
Osmundacese and made the first contribution to the knowledge of the pinna-trace in 
ontogeny. In 1912 Bower described in detail the pinna-traces of Lophosoria pruinata, 
Pr., and Gleichenia linearis (Burm.) Clarke, and placed some weight upon the 
structure of the pinna-trace as a phyletic criterion. In working through the vascular 
anatomy of Peranema cyatheoides, D. Don, I found a difference in the type of 
pinna-trace departure in the basal and terminal pinnae, and, following the lead of 
Chrysler ('10, p. 5), named the one " extramarginal " and the other "marginal" 
(Davie, '12). 

In the marginal type the first indication of the preparation for the departure of 
TRANS. ROY. SOC. ED1N.. VOL. L. PART 11. (NO. 11). 48 



350 MR R, C. DAVIE ON 

the pinna-trace, as the leaf-trace is followed up the petiole, is an extension of the 
mass of tracheides on the adaxial side of the petiole. In Asplenium obtusatum* 
Forst. (PL XXXIII. fig. l), which may serve as an example of this type, the two masses 
of xylem are curved in outline, the convex sides of the curves being directed towards 
each other. The adaxial extremities of these xylem-masses become extended towards 
the adaxial corners of the petiole. As the pinnae are not inserted exactly opposite to 
each other, first the extremity of one and then that of the other xylem-mass is separated 
off from the parent trace. Phloem completely surrounds the xylem in both portions 
of the leaf-trace, and the marginal tracheides when nipped off are surrounded by a 
narrow ring of phloem. There are protoxylem elements at the abaxial tips of the 
leaf-trace and on the adaxial sides, not far from the ends. The pinna-trace has one 
small protoxylem group on one side at the time of separation. 

As the tracheides at the margin of the leaf-trace strand are separated from the 
parent strand to supply the pinna-trace, this type of pinna-supply is termed the 
" marginal " one. 

The marginal type is not always as simple as this, however. In Loxsoma 
Cunninghami, R. Br. (PI. XXXIII. figs. 5 and 6), the leaf-trace in the base of the petiole 
is curved in outline. It has a little hook at each extremity. Below the first pinna 
this trace extends laterally ; the hook is pushed further and further from the centre 
of the petiole, a widening of the leaf-trace takes place in the antero-posterior 
direction just back from the hooked extremity (PI. XXXIII. fig. 5) ; this widened 
region becomes arched like the parent trace, and the arched extremity is nipped off 
as a pinna-trace (PI. XXXIII. fig. 6). 

This is more complicated than the process in Asplenium obtusatum, the presence 
of the hooked extremities dictating the complexity. 

In Balantium culcita (L'Herit.) Klf. (PI. XXXIV. fig. 13) the leaf- trace has small 
terminal hooks. Its outline is at first that of a simple arch with an almost fiat 
abaxial portion flanked by curves (text-fig. l). As the trace ascends towards the first 
pinnae it becomes extended towards the adaxial corners of the petiole ; an arching 
more marked than that of Loxsoma goes on in the extensions, and the pinna-trace 
departs as a replica of the parent trace, taking with it the original margin. 

The simplest example of the marginal type of supply from a hooked leaf-trace is 
found in Aneimia hirta (L.) Sw. (PI. XXXIII. fig. 3). Each hook is merely a little 
group of tracheides placed at right angles to the set composing the side of the leaf- 
trace. The pinna is supplied from an extension of the extremity in the usual manner. 
A swelling in the side of the leaf-trace not far from the tip is the breaking-point for the 
pinna-trace. The new hook of the leaf-trace is the remnant of the dilated portion. 

All four types agree in nipping off the margins of the leaf-traces to supply 
the pinnae. 

* For the sake of uniformity, the nomenclature throughout this paper is that of the Index Filicum (Chbistenskn, 
1806). 



THE PINNA-TRACE IN THE FERNS. 



351 



The " extramarginal " types differ from them in leaving the leaf-trace margins 
intact throughout the petiole. In Didymochlsena truncatula (Sw.) J. Sm. (PI. XXXIII. 
fig. 7), the leaf-trace is composed of several strands. The two adaxial strands have 
their margins sharply recurved, so that they are almost parallel to the adaxial 
surface of the petiole. To supply the basal pinna, the back of the hook is lengthened 
towards the adaxial corner of the petiole. Across the narrow space between the 
two strips of tracheidal tissue thus formed new tracheides spread. The original 
outline of the leaf-trace strand is thus re-formed, while a ring of tracheides is attached 
to the back of its hooked portion. This ring soon separates, to pass into the base of 
the pinna and divide up into a pinna-trace resembling the leaf-trace. The margin of 




Text- Fig. 1. —Balantium culcita (L'Herit. ) Klf. Successive stages in the development and liberation of the pinna-trace. 



the parent trace remains intact during the process. Since the pinna-trace goes 
off from the outside of the parent strand, the method of supply may be termed 
" extramarginal." 

The process of supplying the basal pinnae is exactly the same in leaves with 
unbroken leaf-traces, such as that of Odontosoria chinensis (L.) J. Sm., var. Veitchii 
(PI. XXXIV. fig. 9). 

In this species and in Didymochlxna truncatula no gaps are left in the tissues 
of the parent trace when the pinna-trace is given off. Sometimes, however, the 
departure of the pinna-trace makes a gap in the tissues at the back of the hook in 
the leaf-trace. This is readily seen in Leptopteris hymenophylloides (A. Rich.) Pr. 
(PI. XXXIII. fig. 8). Hymenophyllum demissum (Forst.) Sw. (PI. XXXIV. fig. 10) 
gives off the supply to the basal pinnae in the extramarginal manner. The tracheides 
of the leaf-trace do not form a perfectly continuous series, but the marginal set, 
which terminate the hook of the leaf-trace, remain in position when the pinna-trace 
departs. It can hardly be said that a gap occurs opposite to the departing pinna- 
trace, for a single tracheide occurs between the end of the abaxial curve and the 



352 MR R. C. DAVIK ON 

marginal set of tracheides. The method of pinna-supply in Blechnum orientals, L. 
(PL XXXIV. fig. 11), is the most nearly marginal among the extramarginal types. 
Only one or two of the marginal tracheides remain when the pinna-trace departs, but 
the method of separation of the pinna-trace is certainly extramarginal. The typical 
lengthening of the hook, the bridging of the narrow space between the adaxial curve 
and the margin of the hook, and the retention of the marginal tracheides are all 
found in the process in this Fern. 

' Thin partitions " divide the bounds of marginal and extramarginal types. Gymno- 
gramma Pearcei, Moore, var. robusta (PL XXXIV. fig. 1 2), Ceropteris calomelanos 
(L.) Unci., and C. calomelanos, var. chrysophylla, Klf., have the same type of leaf-trace 
strand as Blechnum orientate. The pinna-trace goes off from a lengthened leaf-trace 
strand, which, however, re-forms its hooked extremity from a swelling of tracheides 
back from the margin. This swelling never catches up the marginal set of tracheides 
of the original leaf-trace. The Gymnogramma type is but a step from the type 
described in Blechnum orientale. 

Histiopteris incisa (Thbg.) J. Sm. (PL XXXIV. fig. 15 and PL XXXV. fig. 16) illus- 
trates a curious combination of the two types in its process of pinna-supply. The 
leaf-trace has the outline of that of Balantium culcita — a flat-topped arch with in- 
curved sides. Small hooks are present at its extremities. The hooked extremity 
lengthens in the usual way and gives off its tip in marginal fashion. Simultaneously 
the projecting corner of the arch beside it also lengthens, and from it is nipped off a 
ring of vascular tissue in the extramarginal manner (PL XXXIV. fig. 15). The two 
strands pass towards the base of the pinna, come together, and by the opening out 
of the adaxial side of the extramarginally-derived portion form a pinna-trace 
exactly like the leaf-trace (PL XXXV. fig. 16). 

In one or two broken leaf-traces a reminiscence of this combined process seems to 
appear. All three strands are unhooked in Aspidium Moorei (Hk.) Diels. Part of 
the pinna-trace comes from the tip of the adaxial strand, part from the median strand 
— those parts which correspond to tip and arch-corner in Histiopteris incisa. The 
pinna-trace of Leptocliilus cuspidatus (Pr.) C. Chr. also comes from portions of the 
leaf-trace corresponding to those supplying the pinna in Histiopteris incisa. The 
leaf-trace is made up of six or seven strands (text-fig. 2). The adaxial pair have 
incurved extremities and give off" the backs of their hooks to supply part of the 
pinna-trace. The rest of the pinna-trace comes from the two subsidiary strands next 
in order to the adaxial strand as we go towards the abaxial side of the petiole. These 
subsidiary strands have simple plates of xylem, from which the strands going to the 
pinnae are simply nipped off. 

The pinna-traces of most of the Cyatheacese bear a resemblance to these. 
Gwynne-Vatjghan ('03) has called attention to the distinction made by Bertrand 
and Cornaille ('02) between the two regions of the petiolar trace, (l) the abaxial 
curve, and (2) the adaxial arcs. The pinna-trace of Cyathea Brunonis comes partly 



THE PINNA-TRACE IN THE FERNS. 



353 



" from the point where the adaxial arc or the xylem hook joins on to the abaxial 
curve," and partly " from the abaxial curve itself at the point where it is folded 
inwards." These are just the points of departure of the pinna-trace in Histiopteris 
incisa and Leptochilus cuspidatus, but the involution of the trace in Cyathea 
Brunonis and its folding to the abaxial curve make both strands in it go off extra- 
marginally. 






Text-Fig. 2. — Leptochilus cuspidatus (Pr.) C. Chr., showing the method of pinna-supply from the leaf-trace. 



These types of pinna-trace cover the variations which are found in the genera 
examined. Upwards of a hundred and fifty species have been worked through, and 
the method of vascular supply to all the pinnae noted from base to apex of the leaf. 
The species were at first chosen at random, but soon a general trend could be observed 
through the families in phyletic order. And the later species examined were selected 
because of their positions on phyletic lines. The classification adopted in the 
accompanying table of the results (p. 354) is that of Diels in Engler and Prantl's 
NaLiirliche Pflanzenfamilien, a classification which corresponds closely to Bower's 
grouping into Simplices — Gradatee — Mixtse. 

Starting with the Leptosporangiate Ferns, we have the Osmundacese showing 
extramarginal supply to the pinna-traces. The length of the gap may vary. It is 



354 



MR R. C. DAVIE ON 



Summary of Result* of Investigation. 



Family. 



Si-hizivaceae 



Osmundaceae 



Gleicheniacese 



Hymenophyl 
laceae 



Loxsomacea? 



Cyatheaceoe 



Woodsieae 



Aspidieae 



Davalliete 



Asplenieae 



Pterideae 



Polypodieae 



Marginal Type of Pinna-Supply. 



Extramarginal Type of Pinna-Supply. 



Lygodium circinnatinn (Burm.) Sw. ; L. scandens (L.) Sw. 
Mohria eaffrorum(L.) Desv. 

Aneimia collina, Raddi ; A. hirta (L. ) Sw. ; A. phyllitidis 
(L ) Sw. ; A. rotundifolia, Schrad. 







... 


Loxsoma 


Cun.ningh.auvi, 


R. Br. 



Balantium culeita (L'Herit.) Klf. 
Dicksonia fibrosa, Col. 



Cystopteris fragilis (L.) Bernh.; 0. montana (Lam.) Bernh. 



Nephrolepis Amerpohlii, hort.; N. Fosteri, hort., Hill; N. 

Piersoni, hort. ; N. Scottii, hort. 
Humata repens (L. fil.) Diels, var. alpina. 
Microlepia hirsuta (J. Sm. ) Pr. 
Lindsaya repens (Bory) Bedd. 
Odontosoria retusa (Cav.) J. Sm. 
Davallia assamica (Bedd.) Bak. ; D. bullata, Wall.; D. dis- 

seeta. J. Sm. ; D. immersa, Wall.; D. pallida, Mett. ; D. 

pentaphylla, Bl. ; D. solida (Forst.) Sw. ; D. solida, var. 

tijiensis. 



Asplenium adiantum nigrum, L. ; A. bulbiferum, Forst., var. 

Fabianum ; A. bulbiferum, var. Hillii ; A. obtusatum, Forst. ; 

A. prsemorsum, Sw. ; A. ruta-muraria, L. ; A. tenerum, Forst. 
Ceterach offieiuaruni, DC. 



Gymnogramma Pearcei, Moore, var. robusta. 

Cerojiteris calomelanos (L.) Und. ; C. calomelanos, var. chrvso- 

phylla, Klf. 
Pellaa hastata (Tlibg.) Prantl ; P. rotundifolia (Forst.) Hk. 
Cryptogramma erispa (L. ) R. Br. 
• ]i<-ilanthesargentea(Gmel.) Kze. ; C. myriophylla, Desv., var. 

elegana 
Notliol-ena affinis (Mett. ) Moore ; N. bonariensis (Willd.) C. Chr. 
l'l.iis biamita, L. ; P. cretica, L. ; P. umbrosa, R.Br. 



Poly podium aurcurn, L. ; P. brasiliense, Poir.; P.phymatodes, 

L.; P. Schneideri, liort. , Veitch ; I', vulgare, L. 
Drynaria rigidula (Sw.) Bedd. 



Todea barbara (L. ) Moore. 

Leptopteris hymenophylloides (A. Rich.) Pr. 

Osmunda javanica, Bl. ; 0. regalis, L. 



Gleiehenia circinnata, Sw. , var. speluncae ; G. flabellata, R.Br.; 
G. rupestris, R.Br. 

Hvmenophyllum demissum (Forst. ) Sw. ; H. dilaiatuni (Forst.) 

Sw. 
Trichomanes elegans, Rich.; T. radicans, Sw. 



Cibotium Schiedei, Schlecht. et Cham. 

Thyrsopteris elegans, Kze. 

Cyathea mexicana, Schlecht. et Cham. ; C. pubescens, Mett. 

Hemitelia grandifolia (Willd.) Spr. 

Alsophila glauca ( Bl.) J. Sm. 



Peranema cyatheoides, Don. 

Diacalpe aspidioides, Bl. 

Woodsia ilvensis (L. ) R.Br.; W. polystichoides, Eat. 

Matteucia orientalis (Hk.) Trev. ; M. struthiopteris(L.) Todaro, 

Onoclea sensibilis, L. 



Dryopteris filix mas (L. ) Schott ; D. phegopteris (L.) C. Chr.; 

D. pulvinulifera (Bedd.) 0. Ktze. ; D. serrata (Cav.) C. Chr.; 

D. setigera (Bl. ) 0. Ktze. 
Didymochlaena truncatula (Sw. ) J. Sm. 
Polystichum aculeatum (L. ) Schott., var. angulare, Pr. ; P. 

Standishii (Moore) C. Chr. 



Leptolepia novae-zelandiae (Col.) Kuhn. 

Microlepia hirta (Klf.) Pr. ; M. hirta, var. cristata ; M. platy- 

phylla (Don), J. Sm. ; M. speluncae (L. ) Moore ; M, 

(Thbg.)Pr. 
Dennstaedtia adiantoides (H. B. Willd.) Moore. 
Odontosoria chinensis (L.) J. Sm. , var. Veitchii. 



Blechnum attenuatum (Sw.) Mett.; B. brasilienso, Desv.; B. 
capense (L. ) Schlecht. ; B. discolor (Forst.) Kevs. ; B. Moorei, 
C. Chr.; B. tabulare (Thbg.) Kuhn ; B. Baiiksii (Hk. fil.) 
Mett.; B. laneeolatum (R.Br.) Sturm ; B. Patersoui (li.Br.) 
Mett. ; B. occidentale, L. ; B. orientale, L. 

Doodia asj)era, R.Br., var. multifida. 

Brainea insignis (Hk.) J. Sm. 

Athyrium filix femina (L. ) Roth. 

Diplazium celtidifolium, Kze.; D. marginatum (L.) Diels. 

Woodwardia radicans (L. ) Sm. 



Trismeria trifoliata (L. ) Diels. 

Adiantum polyphyllum, Willd.; A. sanctse catherinse, hort, 

J. Sm. 
Notholii'na sinuata (Lag.) Klf. 



THE PINNA-TRACE IN THE FERNS. 



355 



long in Leptopteris hymenophylloides and in Osmunda javanica, Bl., quite short in 
O. regalis, L. The pinna-traces of Leptopteris hymenophylloides and Osmunda 
regalis are wide ; those of Todea barbara (L.) Moore and 0. javanica are narrow. 

The Schizaeaceae, in contrast, invariably supply their pinnae in the marginal 
fashion. In Lygodium circinnatum (Burm.) Sw. and L. scandens (L.) Sw., the leaf- 
trace is unincurved and the supply is quite directly marginal. It is so also in Mohria 
caffrorum (L.) Desv. The hooked leaf-trace already described for Aneimia hirta is 
found also, though less prominently, in Aneimia collina, Raddi (PI. XXXIII. fig. 4), 
while slight hooks also appear in A. phyllitidis (L.) Sw. and A. rotundifolia, Schrad. 

In all of these the pinna-trace goes off marginally. 

The third family of the Simplices — the Gleicheniaceae — is consistently extra- 
marginal in supplying its pinnae. Boodle ('01) and Bower ('12) have described the 







Text-Fig. 3. — Gleichenia Jiabellata, R. Br. Diagrammatic representation of the changes occurring in the 
vascular system of the petiole during the liberation of the pinna-traces. 



pinna-traces of Gleichenia dicarpa, R. Br., var. longipinnata, G. Jiabellata, R. Br., and 
G. linearis (Burm.) Clarke. In Gleichenia Jiabellata (PI. XXXIV. fig. 14) the incurved 
margins become nipped off from the rest of the leaf-trace on its inner side, round 
themselves into little rings of tracheides, and run up isolated from the main part of the 
trace for some distance. The back of the remaining hook separates off to go to the 
pinna, and then these isolated circles open out on their abaxial sides and re-form the 
leaf -trace by extending to meet its margins (text-fig. 3). In Gleichenia rwpestris, 
R. Br., the margins of the leaf-trace separate off when the pinna-traces are being- 
given off, but here they do not form two rings but only a couple of arcs with the 
concave sides towards the centre of the petiole. They may even become attached 
to each other by their margins. In Gleichenia circinnata, Sw., var. speluncse, the 
pinna-supply is of the normal extramarginal type ; almost no gap occurs in the 
tissues of the leaf-trace when the pinna-trace is given off. 

The method of supply in Gleichenia Jiabellata and G. rupestris may be regarded 
as a special variety of the type seen in the Osmundaceae, as long gaps are thus formed 
in the leaf-trace opposite to each pinna. 

Passing to the Gradatae, we find the extramarginal type prevails. In the 



356 MR R. C. DAVIE ON 

Hymenophyllacese it occurs in Hymenophyllum demissum, H. dilatatum (Forst.) 
Sw., Trichomanes elegans, Rich., and T. radicans, Sw. 

A divergence into the marginal type occurs in Loxsoma Cunning hami, described 
above, while the type of Balantium culcita is repeated among the Cyatheaceae in 
Dicksonia fibrosa, Col. In the other members of the Cyatheaceae examined the 
extramarginal type prevails. These include Cibotium Schiedei, Schlecht. et Cham., 
Thyrsopteris elegans, Kze., Cyathea mexicana, Schlecht. et Cham., C pubescens, 
Mett., Hemitelia grandifolia (Willd.) Spr., and Alsophila glauca (Bl.) J. Sm. 

Passing into the Ferns on the border-line between Gradatse and Mixtae, we have 
the broken leaf-trace in Peranema cyatheoides, Don, and Diacalpe aspidioides, BL, 
the unbroken leaf-trace in Woodsia ilvensis (L.) P. Br., W. polystichoides, Eat., 
Matteucia orientalis (Hk.) Trev., M. strutliiopteris (L.) Todaro, and Onoclea sensi- 
bilis, L. (in the last three the leaf-trace is binary at its departure from the stalk, but 
unbroken below the pinnae). In all of these the pinna-trace goes off extramarginally 
from the back of the hook, whether in the unbroken trace or in the adaxial strand of 
the broken type. 

- In the genus Cystopteris there is a variation ; in Cystopteris fragilis (L.) Bernh. 
and C. montana (Lam.) Bernh., the supply to the pinnae is marginal. The leaf-traces 
have slightly hooked ends to each portion of the binary strands. The whole set of 
tracheides lengthens, a "thickening" of the number occurs about the centre of the 
leaf-trace on the adaxial face, a break takes place there in the series, and the 
" daughter " trace separates off. Below the second pair of pinnae the binary trace 
becomes an unbroken strand, and the supply is there distinctly marginal. The supply 
to the basal pinnae resembles the type already described for some species of Gymno- 
gramma and Ceropteris (p. 352), though the thickening is less marked in Cystopteris. 

In the Aspidieae, the leaf-traces are of the broken type and the supply to the 
pinnae is extramarginal. In Dryopteris phegopteris (L.) C. Chr. there is a gap 
opposite to the pinna. In Dryopteris pulninulifera (Bedd.) 0. Ktze., D. filix mas 
(L.) Schott, D. setigera (Bl.) 0. Ktze., D. serrata (Cav.) C. Chr., Didymochlsena 
truncatula, Polystichum aculeatum (L.) Schott, var. angulare, Pr., P. Standishii 
(Moore) C. Chr., the supply is extramarginal. In Aspidium Moorei (see above, p. 352), 
the pinna-trace comes partly, marginally, from the tip of the adaxial strand, and 
partly from the median strand of the leaf-trace. 

From the Aspidieae forwards into the main mixed Ferns both extramarginal and 
marginal types of pinna-supply occur. The Davallieae carry forward the extra- 
marginal type through Leptolepia novsB-zelandise (Col.) Kuhn, Microlepia hirta (Klf.) 
Pr., M. hirta, var. cristata, M. speluncse (L.) Moore, M. platyphylla (Don) J. Sm., 
M. strigosa (Thbg.) Pr., and Dennstaedtia adiantoides (H. B. Willd.) Moore. Micro- 
lepia hirsnta (.J. Sm.) Pr., Nephrolepis Amerpoldii, hort., N. Fosteri, hort., Hill, 
N. Piersoni, hort., and N. Scottii, hort., all distinctly show the marginal type. 
Humata repens (L. fil.) Dicls, var. alpina, and Lindsaya repens (Bory) Bedd., are 



THE PTNNA-TRACE IN THE FEUNS. 357 

also marginal. In the genus Odontosoria both types appear. Odontosoria chinensis, 
var. Veitchii (see above, p. 351), is pronouncedly extramarginal, but 0. retusa (Cav.) 
J. Sm. is marginal. The marginal type occurs in Davallia assamiea (Bedd.) Bak., 
D. bullata, Wall., D. dissecta, J. Sm., D. immersa, Wall., D. pallida, Mett., D. 
pentaphylla, Bl., D. solida (Forst.) Sw., and D. solida, var. Jijiensis. 

This appearance of both types in a number of species within a family is repeated 
again in the Asplenieee ; but in them the extramarginal type is restricted to certain 
genera, the marginal type to others. 

A marked and consistent adherence to the extramarginal type is found in the 
genera Blechnum, Doodia, Woodivardia, and Brainea. In Blechnwn attenuatum 
(Sw.) Mett., B. brasiliense, Desv., B. capense (L.) Schlecht., B. discolor (Forst.) Keys., 
B. Moorei, C. Chr., B. tabulare (Thbg.) Kuhn, Doodia aspera, R. Br., var. multifida, 
and Brainea insignis (Hk.) J. Sm., the type of supply is like that seen in Didy- 
mochlsena truncatula, the pinna-trace departing from the back of a very distinctly 
incurved adaxial leaf-trace bundle. 

An even more pronounced form of the extramarginal type is found in Wood- 
ivardia radicans (L.) Sm., where the large pinna-trace arches up almost before it is 
free from the parent bundle, and then breaks into several separate strands. 

In Blechnum Banksii (Hk. fil.) Mett., B. lanceolatum (R. Br.) Sturm, and 
B. Pater soni (R. Br.) Mett., the adaxial leaf-trace bundles are short and narrow, 
and the number of tracheides in the hooked ends is quite small. The pinna-trace 
is undoubtedly extramarginal, though only a few tracheides of the margin are 
left when it has gone oft'. Blechnum occidentale (L.) is also extramarginal, but 
approaches very closely to B. orientale, which is only just extramarginal. 

The other extramarginal members of the Aspleniese are Athyrium filix femina (L.) 
Roth., Diplazium celtidifolium, Kze., and D. marginatum (L.) Diels, in which the 
large pinna-traces go off from the back of wide leaf-traces with distinct terminal 
hooks. The amount left of the original margin after the leaf-trace has gone off is 
greater in Athyrium filix femina and Diplazium marginatum than in Diplazium 
celtidifolium. 

In the genus Asplenium the supply is consistently marginal. In Asplenium 
bidbiferum, Forst., var. Fabianum, and in A. bulbiferum, var. Hillii, A. adiantum 
nigrum {h.), A. prsemor sum, Sw., A. obtusatum, A. tenerum, Forst., and A. ruta- 
muraria (L.), species which differ from one another in size of leaf, in complexity of 
leaf-segmentation, and in leaf-texture, the supply departs on the marginal plan. 
Ceterach officinarum, DC, also supplies its pinnae from the margin of its leaf-trace. 

This division of a family between the one type and the other occurs also in the 
Pteridese, where some genera show the marginal, some the extramarginal type. Two 
genera show some species following the marginal type and others following the extra- 
marginal. Trismeria trifoliata (L.) Diels and Adiantum sanctse catherinse, hort., J. 

Sm., and A. polyphyllum, Willd., give off their pinna-traces in the normal extramarginal 
TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 11). 49 



358 MR H. C DAVIE ON 

manner. Gymnogramma Pearcei, Ceropteris calomelanos, and C. calomelanos, var. 
chrysophylla, show the marginal type in one of its varietal forms (cf. above, p. 352). 
The marginal type also occurs in Pellsea hastata (Thbg.) Prantl and P. rotundifolia 
(Forst.) Hk., Cryptogramma crispa (L.) R. Br., and in Cheilanthes argentea (Gmel.) 
Kze. and C. myriophylla, Desv., var. elegans (PL XXXV. fig. 21). In these Ferns the 
leaf-trace has a compact mass of tracheides, four-sided and wider on the adaxial than 
on the abaxial side of the petiole. The margin is simply nipped off to supply 
the pinnae. 

This form of leaf- trace appears also in Notholsena affinis (Mett.) Moore and N. 
bonariensis (Willd.) C. Chr., and the pinna-trace departs in the Pellaia- Cheilanthes 
fashion. But in Notholsena sinuata (Lag.) Klf. the leaf-trace is binary, the adaxial 
ends of each strand being distinctly hooked. Here the pinna-trace comes from the 
back of the hook and leaves a fairly long gap in the side of the leaf-trace strand after 
it has left it. 

Two sharply contrasted types of pinna-supply are seen in Pteris and the genera 
associated with it. Pteris cretiea, L., P. biaurita, L., and P. umbrosa, R. Br., have 
an ordinary marginal pinna-supply. But Histiopteris incisa (PI. XXXIV. fig. 15 and 
PI. XXXV. fig. 16) joins Pteris (Litobrochia) macilenta, A. Rich., and Hypolepis 
tenuifolia (Forst.) Bernh. in showing a combination of extramarginal and marginal 
types, the pinna-trace being built up of two parts, one coming extramarginally from 
the corner of the arch of the leaf-trace, the other marginally from its extremity. 

This can only be related to what appears in the Cyatheaceae, in Aspidium Moorei 
and in Leptochilus cuspidatus. It bears no resemblance whatever to the process of 
pinna-supply in other species of Pteris. 

In the Polypodiaceae we have a consistent adherence to the marginal type of 
pinna-supply. Polypodium aureum, L., P. vulgare, L., P. phymatodes, L., P. 
brasiliense, Poir., P. Schneideri, hort., Veitch, and Drynaria rigidula (Sw.) Bedd. 
(PI. XXXV. fig. 20), all have their pinna-traces simply nipped off from the edges of 
the leaf-trace bundles, and never give any suggestion of an extramarginal tendency. 

Turning to the Eusporangiate Ferns, we find that the Marattiacese stand by them- 
selves in their method of giving their pinnae a vascular supply. In Angiopteris 
evecta (Forst.) Hoffm. and in Marattia attenuata, Lab., the pinnae are supplied by 
several separate vascular strands. In the petiole there is a complete ring of vascular 
strands near the periphery, while the central portion is occupied, in Angiopteris* by 
a single wide vascular strand, in Marattia by five or six strands in two groups, four 
or five strands making a curved set towards the abaxial face of the petiole, and one 
being situated below them towards the adaxial face. In both Ferns the pinnae are 
supplied by the strands at the corner of the leaf-trace nearest to the pinna, by one or 
two strands from the abaxial and adaxial edges of the peripheral set, and by one 
strand cut off' from the strand or strands in the central position (text-figs. 4 and 5). 

* In rather small leaves. 



THE PINNA-TRACE IN THE FERNS. 



359 



As the Marattian leaf-trace appears to be the horse-shoe type with much infolded 
margins, the pinnae are supplied partly from the portions which correspond to the 
backs of the hooked portion of the horse-shoe type and partly from the infolded 







Text-Fiu. 4. — Angiopteris evecta (Forst.) Hoffm. Diagrammatic representation of the process of supplying 

the basal pinna with its vascular system. 

margins, which are represented in the Marattian trace by the strands in the central 
position. A gap is left in the leaf-trace for some distance above the pinna. This 
is soon filled up by a movement into it of the leaf-trace strands at its edges and by 
new strands nipped off from these. 





Text-Fig. 5. — Murattia attenuata, Lab. Diagrams illustrating the separation of the basal pinna-trace from 

the vascular strands of the petiole. 

A similar process is seen in the petiole of Helminthostachys zeylanica (L.) Hk. 
There the pinna is supplied by strands which come partly from those forming the 
outer ring, and partly from the internal petiolar strand (cf. G Wynne- Vaughan, 
'05, p. 266). This type of pinna-supply can hardly be called either marginal or 
extramarginal. 

The case of Helminthostachys is not typical for the Ophioglossacese, however, for 
in Botrychium lunaria (L.) Sw. we do have a definitely marginal type of pinna- 



360 MR R. C. DAVIE ON 

supply (PL XXXV. fig. 17). The small plate of tracheides in the leaf-trace of this 
species extends towards the pinna and gives off its margin as the pinna-trace. 

Chrysler has shown that this marginal type is not invariably found in the species 
of Botrychium. Botrychium virginianum nips off the edges of its leaf-trace prior 
to supplying its pinnae, but these edges move along the inner faces of the leaf-trace 
strands and unite with them. The margins of the re-formed leaf-trace strands then 
pass off to the pinnae. 

These excursions outside of the Leptosporangiate Ferns help us but little in the 
interpretation of the great majority of forms of pinna-trace supply. The Marattian 
and Ophioglossean types of pinna-supply are apparently dependent on the special 
features of leaf-trace in these groups, and must be brought into line with the Filical 
types rather than employed as a means for their elucidation. 

Two other regions of investigation may be entered before we attempt to make any 
general statements about the pinna-traces we have examined. There is first the 
supply to the terminal pinnae. In general this has been found to be marginal. No 
matter what the outline of the leaf-trace in the lower part of the petiole, it generally 
becomes reduced to a simple narrow plate of tracheides with adaxial and abaxial 
strips of phloem lining it in the distal portion of the rachis. Often the supply is 
marginal to the pinnae situated third or fourth in position below the tip of the leaf. 
In such pronouncedly extramarginal forms as Histiopteris incisa, Pteris macilenta, 
and Hypolepis tenuifolia the supply is marginal to the terminal pinnae and to one 
or two below them. In H. incisa the supply is marginal to the pinna eighth in 
order from the tip of the leaf. 

In the Cyatheaceae we might expect a retention of a complicated outline for the 
leaf-trace up to near the tip of the leaf. In Cibotium Schiedei the strand is below 
the ultimate pinna an ellipse of tracheides, very " thin " on the adaxial side. The 
pinna is supplied by the departure of the lateral portion of this ellipse. Though the 
leaf-trace is a closed ellipse, the margins are, of course, on the adaxial side, and this 
supply is really of the extramarginal type. Exactly the same kind of trace appears 
below the ultimate pinna in the leaf of Adiantum polyphyllum, where the pinna is 
supplied after the same fashion. 

In the early stages of the life of a Fern the method of pinna-supply is the 
marginal one. In the first pinnae in a very early leaf of a plant of Athy7'ium filix 
fernina the supply came from the edges of a simple plate of tracheides, in exactly 
the manner seen in most terminal pinnae of mature leaves. In a later leaf the trace 
was binary and showed a tendency to form hooks at the edges of the two parts. 
Here the supply to the basal pinnae was again marginal, though not from the extreme 
tip of the trace. 

In a young plant of one of the Cyatheaceae the supply was marginal in the first 
leaf, extramarginal in one considerably older (about the eighth). 

In ;i relatively late leaf of a plant of Peranema cyatheoides (about the seventh 



THE PINNA-TRACE IN THE FERNS. 361 

leaf) the supply to the basal pinnae was marginal, of the type seen in the supply to 
the basal pinnae in Cystopteris montana. 

In a young plant of Asplenium bulbiferum, the supply to the basal pinnae in the 
first leaf was marginal. The xylem formed a narrow plate, its long axis at right 
angles to the radius of the stem of the plant. From the margins of this line of 
tracheides the pinna-trace was simply nipped off. 

In a young plant of Asplenium lunulatum, Sw., sub-sp. erectum, Bory, the supply 
to the basal pinnae in a very early leaf was marginal. In the ninth or tenth leaf the 
supply was also marginal to the basal pinnae. In the earlier leaf the group of 
tracheides was arranged in a compact mass, wide and slightly hollowed on the 
abaxial side and with two diverging extensions towards the adaxial corners. In the 
later leaf the abaxial portion had two little projections, each composed of only a 
few tracheides, while the adaxial extensions were much longer than in the earlier leaf. 
The axis is dictyostelic and the leaf-trace departs from the strands subtending the 
leaf-gap at a point where the two strands almost meet. The leaf-trace is single, 
though made up of two portions of xylem, coming each from one limb of the 
dictyostele. There is a common endodermis round the two portions of xylem and 
their surrounding phloem. 

After the pinna-trace has left the petiolar bundle it develops differently in 
different Ferns. 

Traces like that of Loxsoma are fully formed by the time they pass into the pinna. 

Gleichenia, Osmunda, Todea, Hymenophyllum, Trichomanes, Balantium culcita, 
Dicksonia fibrosa, Histiopteris incisa, Hypolepis tenuifolia, the Cyatheaceae all have 
this type of pinna-trace. 

Ferns with a " broken " leaf- trace (e.g. Diacalpe aspidioides) give off the pinna- 
trace as a " signet-ring," then break it into two in the pinna, and nip off the other 
strands from the aporachial ends of these two. 

In Davallia pallida the pinna-trace passes into the stalk of the pinna as a dorsi- 
ventrally constricted mass of xylem enclosed in phloem (PI. XXXIII. fig. 2). The 
xylem is wider in the centre of the pinna-trace than at the edges, where the margins 
are slightly curved towards the prosrachial side of the stalk. A short distance out 
into the pinna-stalk there appear suddenly in the phloem on the aporachial side, first 
a single tracheide and then a small group, two or three in number (text-fig. 6, b). 
These are quite separate from the main set of tracheides of the pinna-trace, but as 
they increase in number the tracheides of the main trace extend towards them, and 
presently the two sets join (text-fig. 6, c). We have then a pinna-trace of triangular 
outline, the large amount of tracheides— and these greater in size of lumen than the 
others — in the central position, the margins being composed of a narrow set of small 
tracheides. Further out into the pinna the aporachial side of the group of tracheides 
becomes widened out on the peripheral face, so that there are two triangular groups 
attached to each other by the apices of the triangles (text-fig. G, d). A little further 



362 



MR H. C. DAVIE ON 



out still the aporaehial set becomes broken up into two curved portions, and these 
separate from each other at their tips, so that the pinna-trace now has the outline 
of the leaf-trace (text-fig. 6, e). The supply to the pinnules goes off from this in just 
the way in which the pinna-supply leaves the leaf-trace (text-fig. Q,f). But in the 
pinnule there is no development of any accessory group of tracheides, and the pinnule- 
trace retains its narrow plate of tracheides throughout its length, giving off the 
supplies to the secondary pinnules from its margins. 

In Asplenium lucidum, Forst., the supply to the pinna leaves the petiolar trace 
as a narrow strip of tracheides enclosed in phloem. Just as in Davallia pallida, a 
group of tracheides soon appears on the aporaehial side of the pinna-stalk, becomes 
larger, and soon joins the main group of tracheides of the pinna-trace. The triangular 
set of tracheides thus formed soon splits at the apex of the triangle, and two portions, 
each like one part of the binary leaf-trace, remain attached to each other by their 




Text- Fig. 6. — Davallia pallida, Mett. Series of diagrams to show the development of the pinna-trace. 



middles. From this pinna-trace the supply to the pinnules is nipped off* marginally. 
Just as in Davallia pallida, there is no appearance of tracheides in the pinnule-trace 
in the position of their occurrence in the pinna-trace. 

In Drynaria rigidula the pinna-trace leaves the leaf-trace (PL XXXV. fig. 20) 
as a narrow plate of tissue, with about twelve tracheides. The tracheides in the centre 
of the plate are much larger than those at its edges, and as the strand passes out 
into the pinna these central tracheides rearrange themselves, crowding into a little 
four-sided group, the small tracheides remaining as narrow strips at the edges of the 
group. In this condition the vascular strand passes up to the tip of the pinna. 

In Asplenium prasmorsum the margin of the leaf- trace which goes off to the pinna 
remains as a narrow plate right through the pinna, only extending laterally and 
nipping off its edges when pinnules have to be supplied. 

This survey of the pinna-trace and its development and elaboration in the Ferns 
confirms the impression given by an examination of the table on p. 354. There is a 
gradual rise; of the extramarginal type out of the marginal, and then a lessening of 
t In- type until in the most advanced Ferns it disappears altogether and the marginal 
type remains. The comparison of the pinna-traces of the early leaves of various Ferns 



THE PINNA-TRACE IN THE FEENS 363 

shows that at first the marginal type prevails. And an examination of the ultimate 
branches of the Fern leaf reveals the marginal type in the great majority of species. 

We have then to explain how this variation of type has come to be, and whether 
it can be related to any prominent features in the development of the vascular system 
in the Ferns. To what is the variation due ? Can we trace it to the differences in 
the construction of the stele ? Is it dependent on the length of the leaf, on the 
number of pinnae, or on the state of division and subdivision of the pinnae themselves ? 
Or is the outline of the leaf-trace responsible for the differing points of departure of 
vascular supply for the appendages of the leaf ? 

Sinnott has based his classification of leaf-traces upon the number of protoxylem 
groups present in them. On this basis he has separated Osmundaceae from Marat- 
tiaceae, and these from all remaining forms. The Osmundaceae are protagonists of 
the extramarginal type of pinna-supply ; the Marattiaceae are divergent but consistent 
adherents of the same type ; the real differences occur in " all remaining forms." 
The explanation of pinna-trace type cannot be in the number of protoxylem groups 
in the leaf-trace. 

We are brought no nearer to an explanation by the work of Parmentier ('99), # 
whose classification of the Ferns of France on the structure particularly of their 
petioles is not too reliable. For his type-sections of these petioles have apparently 
been made at varying levels, and his arrangement depends much upon the comparison 
of sections taken at different distances from the base of the petiole. No attempt is 
made to compare one leaf-trace with another as they run from end to end of the leaf. 

Two notable contributions to the knowledge of the evolution of the Filicinean leaf- 
trace are those of Thom,e ('86) and Bertrand and Cornaille ('02). In both of these 
papers a comprehensive survey of Fern-petioles is made and distinctive types are 
contrasted with one another. 

From the leaf-traces of Phegopteris, Onoclea, Aspidium, Thom^e makes an 
" A spidium- type," which he joins with the " Asj^lenium-type " and the intermediate 
" Struthiopteris germanica-ty-pe " as a " Polypodiaceous type." 

Against this he sets the " Cyatheaceous type." The transition from the one to 
the other is through Hypolepis tenuifolia,^ Pteris tremula, and Hypolepis repens. 

He makes the Marattiaceae and Cibotium exceptional modifications of these types. 
He concludes that the different families of Ferns can be distinguished by the charac- 
ters of the leaf-trace, but that a systematic arrangement of Ferns on anatomical grounds 
is quite outside of any serious consideration, and that only broad distinctions can be 
confirmed by the anatomy of the petiole. 

The most interesting feature of his classes is the selection of the bridge from the 
Cyatheaceous type to the Polypodiaceous type. For we have seen that the pinna- 

* Cf. PELOURDE ('06). 

t In quoting the work of Thom^e and Bertrand and Cornaille I retain their nomenclature of the Ferns they 
describe. 



304 MR K. C. DAVIE ON 

trace of Hypolepis tenuifolia relates that of the Cyatheacese to that of certain species 
of Pterin. His contrast of an Aspidium- Asplenium type with that of the Cyatheacese 
finds an echo among the pinna-traces. So, too, does his opposition of the Aspidium 
leaf-trace to that of Asplenium. 

A wider and more ambitious treatment of the leaf-trace is given by Bertrand 
and Cornaille. No account of Fern-leaf anatomy can give other than a careful and 
grateful consideration of this exhaustive work. It abounds in curious details of 
minutest observation, many of them tucked away, unfortunately, in little corners 
where they easily lurk unnoticed. The Osmundean trace (that of O. regalis) is 
regarded as the prototype from which those of other Ferns are derived. The traces 
of Todea barbara, T. Fraseri, T. superba, T. hymenophylloides, Aneimia collina, 
A. phyllitidis, Mohria eaffrorum, Gleichenia dichotoma, and G. rupestris are taken 
as variations of this Osmundean trace. The Cyathean trace of Dicksonia antarctica 
and D. squarrosa is the Osmundean trace with refolded adaxial arcs. Its variations 
occur in Matonia pectinata, Dicksonia regalis, Cyathea medullaris, Alsophila 
australis, and Hemitelia Smithii. Then comes the Onoclean trace of Struthiopteris 
germanica. A variant on it in Onoclea sensibilis prepares for Polypodium phego- 
pteris and Lomaria spicant, while it appears, greatly reduced, in Nephrodium molle 
and Doodia aspera. 

Within the wide range of variations of this type come the traces of Davallia 
repens, Scolopendrium officinale (the " polypodian hippocampus"), Davallia fmni- 
culacea, Asplenium ruta-muraria, A. tricliomanes (reduced in Marsilia and Pilularia), 
A. Nidus-avis, Onychium japonicum, Davallia tenuifolia, and Pellea geraniifolia. 
Amplified variations of the same type appear in Gymnogramma tatarea, Blechnum 
brwsiliense, Lithobrochia vespertilionis, Microlepia platyphylla, Polypodium Hera- 
cleum, Ceratopteris thalictroides, Goniopteris proliferum, Dennstaedtia davallioides, 
and Pteris aquilina, while a very much reduced form appears in Lygodium japonicum, 
L. scandens, Tricliomanes radicans, and Hymenophyllum tunbridgense. A Poly- 
botryan trace, isolated in character, occurs in Lomariopsis fraxinifolia. 

In contrast to these types is a Marattian trace of Marattia fraxinea, M. Lancheana, 
Angiopteris evecta, and Kaulfussia sesculifolia. The Ophioglossean trace of Hel- 
minthostachys zeylanica, Botrychium rutaceum, B. virginicum, and Ophioglossum 
vulgatum is derived from the Marattian trace. 

Obvious situations for tourney-fields lie in the announcement of the Osmundean 
trace as the primitive type and in the assortment of heterogeneous leaf-traces under 
the Onoclean banner. 

No early leaf I have examined shows a leaf-trace like that of Osmunda regalis ; 
many show outlines not unlike that of Aneimia collina and Lygodium scandens. 
A in I a classification of Doodia aspera and Blechnum brasiliense alongside of Asplenium 
ruta-muraria and Scolopendrium officinale, with Onoclea sensibilis, Polypodium 
phegopteris, ami Microlepia platyphylla not far off, gives no clue to the widely 



THE PINNA-TRACE IN THE FERNS. 365 

divergent pinna-traces of these species. These Ferns may all rightly be of the 
Onoclean type, but that avails nought to explain why they supply their pinnae in 
such very different ways. 

But the leaf-trace headlines Osmundean, Cyathean, Onoclean, and Marattian stand 
clear and are suggestive. From the Osmundean and Onoclean groups there may be 
drawn types of leaf-trace which agree in supplying their branches on a definite plan 
which is seen in the earliest known Ferns, and which is found in those Fern-leaves 
which occur first in the development and in those parts of Fern-leaves which are 
regarded as primitive. And from the Onoclean group, too, can be taken other types 
which are now known to be high in the scale and which supply their pinnae in a 
manner, so to speak, improved upon that dominant in the Osmundean and Onoclean 
groups. The marginal type of pinna-supply cuts up the Osmundean and Onoclean 
groups when it is applied as a differential factor. And the marginal type is at once 
the most primitive and the most advanced. 

This presence of the marginal type at the bottom and at the top of the evolutionary 
ladder brings out prominently the need for discovering the factors that produced the 
intermediate extrarnarginal type. This is altogether dependent on the presence of 
the incurved hooks at the ends of the leaf-trace. It only appears where they are 
present, though sometimes hooks are present in leaf-traces that supply their pinnae 
marginally. The evolution of the hooked leaf-trace must be the first factor in this 
development of the extrarnarginal type of pinna-trace. 

This evolution is suggested by Kidston and Gwynne-Vaughan ('08) in their 
description of the leaf-trace of Thamnopteris Schlechtendalii, Eichw. (text-fig. 7). 
There the leaf-trace arises as a swelling of the xylem of the stem. It passes off 
without leaving any gap or depression in the stem-xylem, then protoxylem appears 
almost centrally in it, an island of parenchyma occurs on the adaxial side of the 
protoxylem, this island increases in size, the protoxylem groups widen along the 
bay and the curved trace is produced, followed by a curving of the petiolar outline. 
These changes in the individual leaf-trace are held to indicate the changes under- 
gone in the ontogeny and phylogeny of the adaxially curved leaf-trace so repre- 
sentative of the Filicales. The adaxial hooks thus remain as part of the adaxial 
side of an originally solid leaf-trace. 

Such a solid leaf-trace does occur in several of the Zygopterideae. 

In Dineuron ellipticum, Kidston, and D. pteroides, Renault, the leaf-trace has 
an elliptical strand of xylem grooved at the ends. The pinna-traces depart from 
these ends as curved bars leaving no gaps in the tissues of the leaf-trace (text-fig. 8, a). 
In Metaclep>sydropsis duplex, Williamson, the solid mass of tracheides in the leaf- 
trace has the outline of an hour-glass. The pinna-traces go off from the dilated ends 
of this leaf-trace (text-fig. 8, c). Diplolabis Romeri (Solms) supplies its pinnae from 
the same relative position of the leaf-trace as do the two preceding species, but the 
arms of the leaf-trace are highly developed here in the normal petiole, though at its 
TRANS. ROY. SOC. ED1N., VOL. L. PART II. (NO. 11). 50 



366 



MR R. C. DAVIE ON 



base stages like the leaf-traces of Dineuron and Metaclepsydropsis have been found. 
Etapteris diupsilon, Williamson, has the ends of the arms very much developed, 




Text-Fig. 7. — Diagrams illustrating the departure of the leaf-trace in Thamnoptcris SehleeMendalii, 
Eiohw., sp. (After Gwynne-Vauuhan and Kidstok.) 

though the pinna-trace conns from the middle of the arm (of. text-fig. 8, />), just 
from the position from which the pinna-trace goes off in Dineuron. Unfortunately 



THE PINNA -TRACE IN THE FERNS. 367 

for the sake of minute comparisons with the living Ferns, these Zygopteridese seem 
to have possessed four orthostichies of primary pinnse. The pinna-traces which we 
have mentioned presently divide into two parts each, so that these Ferns must not be 
examined too critically for prototypes of leaf-trace. But there is at least a constancy 
in the position of origin of the pinna-trace-bars, as Gordon terms them. They come 
from the margins of the solid leaf-traces just from the points nearest to the appendages 
which it is their work to supply. 

Kidston and Gwynne-Vaughan have classified the Zygopteridese according to 
the presence of a single row of appendages ("pinnse") on each side of the main 
rachis or of two such rows. 

Those genera already mentioned fall into their second class. The first class, with 
a single row of appendages on each side of the main rachis, includes Ankyropteris 
bibractensis, Renault, and Clepsydropsis antiqua, Unger. From the edge of the 
leaf-trace in Clepsydropsis antiqua the pinna-trace goes off as a closed ring. This 





TEXT-Fro. 8. — Diagrams illustrating the pinna-trace departure in a, Dincuron ptcroidcs, Renault ; 6, EtaplAris Scotti, 
P. Bertrand ; c, Metaclepsydropsis duplex, Williamson. (After P. Bertrand, and Gwynne-Vaughan and Kidston.) 

type of pinna-trace occurs also in Ankyropteris. Ankyropteris corrugata, Williamson, 
has biseriate primary pinnse and one plane of symmetry. The leaf-trace has the 
outline of a double anchor — a modification of the Etapteris-Diplolabis type. The 
pinna-traces in this Fern leave the edges of the arms of the leaf-trace in very much 
the same way as those of Diplolabis leave its leaf-trace, but here there is only the one 
set of pinnse to be supplied from each arm of the leaf-trace, and thus only one half of 
the leaf- trace arm is concerned in the process. It is just a Diplolabis type with half 
of the pinna-trace-bar suppressed. This Ankyropteris-tyye is another illustration of 
the effect of the position of the pinnse relatively to the leaf-trace in affecting the 
point of departure of the pinna-trace. But the details of the departure of the pinna- 
trace in Ankyropteris recall the process in such an extramarginal type as Didy- 
mochlsena truncatula. The full growth of the pinna-trace has not been followed, but 
the sections of Ankyropteris bibractensis, var. westphalensis, P. Bertrand, which 
have been studied by Paul Bertrand (text-fig. 9) (Etudes sur la fronde des Zygo- 
pteridees, Lille, 1909), show the widening out of the tracheides at the part of the leaf- 
trace which is to supply the pinna, the appearance of a reparatory set (" anneau 
reparatrice interne") to ensure the continuity of the tissues of the arms of the leaf- 
trace past the point of pinna-trace departure, and the departure of the pinna-trace 



368 MR R. C. DAVIE ON 

as a closed ring. In Clepsydropsis the actual margin of the leaf-trace goes off to 
supply a pinna ; in Anhyropteris the pinna-trace leaves the leaf-trace in the extra- 
marginal way. 

Where there are two orthostichies of primary pinnae to be supplied from each arm of 
the leaf-trace, as there are in Dineuron, Metaclepsydropsis, Diplolabis, and Etapteris, 
the pinna-trace-bar leaves the leaf-trace on the side directed towards these pinnae 
as a single curved mass of xylem, and then breaks into two parts, each of which moves 




ANNCAU 

REPARATRICE 

INTERNE 




PINNA I- -PINNA 

TRACE /A TRACE 



Text-Fig. 9. — Departure of pinna-trace in Anl-yropteris bibraclensis, var. westphalensis, P. Bertrand. 

(After P. Bkktrand. ) 

out to a pinna. Where the pinnae are uniseriate, as in Anhyropteris, the pinna-traces 
depart from the edges of the leaf-trace at the ends nearest to the pinnae and move 
directly out to them. In Tubicaulis solenites, Cotta (text-fig. 10, a) (the C faces 
with its curve away from the axis — reverse of the living Fern), and in Anachoropteris 
Decaisnei, B. Renault (text-fig. 10, b), the pinna-traces leave the leaf-traces back 
from the edges, and the marginal set of tracheides is undisturbed throughout the 
rachis. But the pinna-trace in these two is a solid patch of xylem and not a closed 
ring. Bertrand makes a suggestive explanation of the Anachoropteris type, deriv- 




o 
o 



O 



Text-Fig. 10. — Diagrams illustrating the departure of the pinna-trace in a, Tubicaulis solenites, Cotta ; 
b, Anachoropteris Decaisnei, Renault. (Alter P. Beutrand.) 

ing it from the type of Dineuron pteroides by development of the anterior hooks. 
He indicates how the position from which the pinna-trace departs in Anachoropteris 
corresponds with that from which the Dineuron pinna-trace goes off from the leaf- 
trace. This is an emphasis of the retention by the pinna-trace of the position of 
departure relatively to the leaf- trace. The pinna-traces of Dineuron and Anaclio- 
ropteris correspond to the marginal and extramarginal types in the living Ferns, 
though we must remember that the Dineuron pinna-trace breaks into two parts after 
Lining off from the leaf-trace, and that it has to supply two orthostichies of pinnae. 
This caution also applies to the pinna-trace of Tubicaulis, though the trace of 
Anachoropteris supplies but one pinna. Any discussion of the type of leaf in the 
Zygopteridese and Botryopteridese as illustrated by these leaf-traces is out of place 



THE PINNA-TRACE IN THE FERNS. 369 

here. The useful information which these Ferns give us is that their pinna-traces 
departed from their leaf-traces at the points nearest to the pinnae. 

If we cany this into the living Ferns, any fundamental distinction between marginal 
and extramarginal types of pinna-trace breaks down — the pinna-traces go off just 
from the points which best serve the pinnse. The distinction between the marginal 
and extramarginal types of pinna-trace must then be one due to the differences in 
leaf-traces, not in the pinnae themselves. This is so far true. But a marked 
divergence from the type general within a family, such as that of Balantium culcita 
(pp. 350, 356) from the type of the Cyatheaceae, must make us pause in the generalisation. 
We have in B. culcita a marginal type of pinna-supply, yet the general type in the 
Cyatheacese is a very elaborate extramarginal one. The changes in the leaf-trace 
of Gleichenia jiabellata throw some light on the interpretation of this divergence. 
There the incurved margins of the leaf-trace round themselves off as circular 
groups of tracheides which run up along the face of the gaps caused by the departure 
of the pinna-traces (PL XXXIV. fig. 14), and then beyond the pinna-trace gaps unite 
to the remaining tracheides of the median part of the leaf-trace (text-fig. 3). The 
incurved edges of the leaf-trace in Gleichenia carry on the water-supply up the 
adaxial face of the leaf. The pinnae are relatively large, and the pinna-traces make a 
considerable drain on the tracheides of the leaf-trace. Apparently such a drain would 
cause a serious deflection of water from the adaxial face of the petiole, and would 
probably cause a water-starvation of the successive sets of pinnae above the basal 
pair. The incurved hooks of the Gleichenia leaf-trace have thus the task of carrying 
forward the water-supply from one pinna-gap to the next, and of providing sufficient 
water to counteract the drain of several sets of pinnae. 

We would thus expect to find a considerable development of incurved hooks in 
leaves possessing many pinnae, and perhaps most distinctly in those leaves with many 
pinnae arranged in close succession (cf. Tansley, Ev. of Fil. Vase. System, p. 117, cf. 
p. 126). In the Cyatheaceae the leaf generally possesses a long "tail" of lamina 
beyond the last pair of actual pinnae. And in the Cyatheaceae the leaf-trace beyond 
the last pair of pinnae is an ellipse of tracheides, thin on the adaxial face (cf. p. 360). 
Just the same arrangement in the leaf-trace is found in Adiantum polyphyllum, 
where the leaf ends in a long " tail " similar to that of Cyatheaceous leaf. Balantium 
culcita has a much shorter leaf than the majority of the Cyatheaceae ; the pinnae are 
relatively not heavy and are arranged at relatively considerable distances from one 
another. Almost the same remarks apply to the leaf of Dichsonia fibrosa, which has 
a marginal pinna-supply just like that of Balantium culcita. These are simple 
Cyatheaceae (perhaps owing their simplicity to reduction) which have modified their 
type of vascular supply to the pinnae in relation to their relatively short leaves. In 
the leaf of the majority of the Cyatheaceae the complicated leaf-trace with its very 
much incurved margins is related to the large size of the leaf, the weight of the 
pinnae, and their large number. The incurved edges of the leaf-trace persist beyond 



370 



MR R, C. DAVIE ON 



even the last pair of pinnae, in order to provide sufficient water-supply for the " tail " 
of the leaf. # The two exceptions, Balantiwm culcita and Dicksonia fibrosa, have 
relatively short leaves with pinnse situated at greater distances from one another 
than are those of Cyathea, Alsophila, and Hemitelia. And these two have only 
slight hooks on the adaxial faces of their leaf-traces, a much simpler type of leaf-trace 
and marginal supply to the pinnse. 

An interesting confirmation of this view of the function of the incurved margins 
of the leaf-trace comes from Botrychium virginianum (Chrysler, '10). There the 




Ticxr- Fig. 1 1. — Botrychium daucifolium, Wall. Diagrams illustrating the departure of the vascular supply 

to the first pair of sterile pinnse. 



margins of the leaf- trace strands nip off, pass along the adaxial face for a short 
distance, and then take up the position of margins for the leaf-trace when the pinna- 
trace goes off. This ensures a continuity of water-supply past the bases of the 
pinnae. Botrychium daucifolium, Wall., affords us an example of a similar process 
(text-tig. 1 I ). In the preparation for the departure of the vascular supply to the 
first sterile segments of the leaf there appears on the inner side of one of the two 
lea I'-t race strands at first a single tracheide, then a small group of tracheides (text- 
fig. I I, b), which approaches the leaf-trace strand and finally unites with it (text- 
fig. I I, <■). Part of the group then goes off with the portion of the leaf-trace strand 

* See also Diplazium marginatum, where the supply is marginal to the nerve fourth in position from the ti j> 
of the leaf. 



THE PINNA-TRACE IN THE FERNS. 371 

going to the pinna, the rest remains as the new margin of the leaf-trace strand 
(text-fig. 11, d, e). This process appears in the supply of the second pinna (text- 
fig. 11, e,f, g), but in relation to the other of the two leaf-trace strands.* 

Still other examples illustrating the same process may be cited in Matonia 
pectinata (Seward, '99) and Archangiopteris Henryi, Christ et Gies. (Gwynne- 
Vatjghan, '05). In Matonia the incurved adaxial edges of the leaf-trace unite, 
their margins extend laterally and pass with the lateral portions of the leaf-trace 
into the pinnae (text-fig. 12). The pinnae are thus supplied with an internal system 
which probably meets the demand for a considerable water-supply without unduly 






O 

Text- Fig. 12. — Diagram illustrating the division of the single petiolar stele into the 
vascular strands of the pinnae. (After Seward. ) 

depleting the supply for the rest of the leaf. For the actual backs of the incurved 
hooks remain intact and pass up the rachis to carry water past the point of pinna- 
trace departure. This looks like a special case of the Gleichenia-type in which not 
only must water be carried up past the pinnae for the needs of the higher parts of 
the leaf, but where the pinnae must also be provided in quick succession with a good 
water-supply. 

The pinna of Archangiopteris Henryi is supplied by strands which occupy much 

* Chrysler argues from similar processes in B. ternatum that we have in this reinforcement a relic of an earlier 
system of passage of the margin along the face of the leaf-trace strand, such as we have already mentioned in B. 
virginianum. He sees no possible use for the reinforcing strand, as it is not connected at its base with the leaf-trace, 
and declares that it cannot convey water. But the structure of the tracheides in the Ferns is such that water might 
quite well be conveyed laterally from tracheide to tracheide even where the sets of tracheides are not continuous at 
their bases. Water can readily be deflected from the main strand into this reinforcing strand as soon as the two are 
in lateral contact, and thus the effect of the drain on the water-supply by the pinna-departure may be compensated for. 



372 MR K. C. DAVIK ON 

llic same position in the rachis as do the parts of the leaf-trace which supply the 
pinna of Matonia pectinata. In Archangiopteris, however, the details are a little 
different, since the leaf-trace has a different outline from that of Matonia. One 
strand which goes into the pinna comes from the side of the curve of the leaf-trace 
set; the other comes from the abaxial face of the adaxial strand (text-fig. 13). The 
terminal part of the adaxial strand remains to carry the water past this pinna-gap. 
This terminal strand is itself reinforced below the departure-point of the pinnae by 
internal accessory strands which come from the internal face of the abaxial curve. 

Wherever we find hooked adaxial portions in a leaf-trace, we do not find any 
evidence of a process of reinforcement, prior to the departure of the pinna-traces, of 
the parts of the leaf-trace supplying the pinnae. But the incurved hooks do vary 
considerably in length and in their number of tracheides, according to whether 
the leaf is a long one or a short one, one with heavy pinnae or one with light pinnae, 
one with closely set pinnae or one with pinnae at considerable distances from one 
another. 




PINNA TRACE , 



Text-Fig. 13. — Archangiopteris Henry i, Christ et Gies. Diagram illustrating the 
method of departure of the pinna- trace. (After Gwynne-Vauuhan.) 

An excellent example of the type of hook found in a long leaf with heavy pinnae 
is that of Didymochlsena truncatula (PI. XXXIII. fig. 7). Species of Dryopteris with 
shorter leaves and lighter pinnae, like D. filix mas, have much smaller adaxial hooks. 
In Microlepia hirta and M. platyphytta, where the pinnae are spread over approxi- 
mately the same length of leaf, the adaxial hooks are of almost the same size ; in 
M. speluncse, whose leaves are usually larger, the adaxial hook is larger than in either 
of these. Leaves with relatively heavy pinnae, like Diplazium celtidifolium or D. 
marginatum, have strongly developed adaxial hooks ; those with fairly light pinnae, 
like the Blechnums, have weak adaxial hooks. In all of these the adaxial hook 
appears to cany the water on from pinna-trace to pinna-trace along the leaf quite 
satisfactorily. Whenever the adaxial hook disappears and we get a marginal supply 
to the pinnae, the system which carried forward the water disappears. On the whole, 
the appearance of the marginal type of pinna-supply seems to coincide with a reduc- 
tion in the size or complexity of the leaf. This is by no means invariable, but a 
contrast of such outstanding genera as Athyrium and Diplazium (extramarginal) 
wit I) Asplenium and Ceterach (marginal), or of Ilypolepis and Adiantum (extra- 
marginal) with Pellasa, Cheilanthes, and Cryptogramma (marginal), shows that there 
i- 30me truth in the statement. A glance over the table on p. 354 confirms this 



THE PINNA-TRACE IN THE FERNS. 



373 



opinion that the extramarginal type of supply is found on the whole among large- 
leaved Ferns, the marginal type among the smaller-leaved genera. A few examples, 
like Poly podium aureum (marginal) and Notholsena sinuata (extramarginal), can be 
taken, which, if they stood alone, would prove the contrary proposition. The general 
survey tends to reveal the extramarginal type among large-leaved, heavily pinnate 
forms, the marginal type among the smaller-leaved, more lightly pinnate forms. 
Davallia solida gives us an interesting example of a process which may help the 
elucidation of the adoption of the marginal type of pinna-trace. The leaf-trace has 











h 



Text-Fig. 14. — Davallia solida (Forst. ) Sw. Series of diagrams illustrating the changes in the leaf-trace during the period 

of supplying the lowest pair of pinnae. 

two fairly large adaxial strands, with unincurved margins, and a small median strand 
(text-fig. 14, a). The adaxial strand gives off a portion marginally on the side directed 
towards the lowest pinna (text-fig. 14, b and c). This passes out into the base of the 
pinna ; then another marginal portion is given off from the adaxial strand and follows 
the first strand into the pinna (text-fig. 14, k). The pinna-trace is binary and gives 
off two strands to the first pinnule. During the supply of the pinna by the leaf- 
trace the median strand nips off a small strand while one of the adaxial strands is 
preparing to give off its margin (text-fig. 14, b). This little strand passes to the 
adaxial strand and fuses with it between the giving-off of the first pinna-trace-strand 
and the second (text-fig. 14, h). The median strand itself moves towards the other 
adaxial strand and fuses with it just before it nips off the first strand for the pinna 

on that side (text-fig. 14, e and f). Some of its tracheides remain as the abaxial 
TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 11). 51 



374 MR E. C. DAVIE ON 

portion of this adaxial strand ; the rest simply separate away again and resume their 
position as the median strand (text-fig. 14, g and h). Thus both adaxial strands are 
reinforced by tracheides from the median strand during the period of pinna-trace 
departure. This process may well be compared with that which we have already 
considered in the supply of the pinnules by the pinna-traces of Davallia pallida and 
Asplenium lucidwn, where the pinna-trace was reinforced by a set of tracheides 
appearing on the aporachial side before the pinnules were supplied. It is noticeable 
that, while the leaf-traces which supply their pinnae on the marginal system are 
simpler in outline than those supplying their pinnae extramarginally, there is a 
development usually of hooks on the abaxial ends of the leaf-trace strands {e.g. 
Davallia, PI. XXXIII. fig. 2 ; Gymnogramma Pearcei, var. robusta, PI. XXXIV. 
fig. 12). These probably serve the same purpose as the reinforcing strands of the 
leaf-trace of Davallia solida, or of the pinna-traces of Davallia pallida and Asplenium 
lucidum — they carry on the water-supply past the pinnae, and prevent a too great 
reduction of the water-supply as we approach towards the tip of the leaf. 

These explanations of the outlines of various leaf-traces are by no means provable 
in every leaf-trace which may be considered. But when they are taken in conjunction 
with one another and in relation to certain distinctive types of Fern-leaf they afford 
us the opportunity for a broad generalisation which finds confirmation in the arrange- 
ment of the species of Ferns in the table on p. 354. 

But we have still to consider the possible advantage to be derived from the 
change from the extramarginal to the marginal type of pinna-supply. It seems to 
have occurred in a passage from a larger-leaved to a smaller-leaved form of the Fern 
(cf. above, p. 373). On that basis its occurrence is fairly easily to be explained. For 
in the small-leaved form the adaxial portion of the leaf-trace would be small and 
thin. To supply pinnae from the back of a thin " hook would be to give them a 
very exiguous pinna-trace. A long strip of tracheides can be more easily given off 
from an unincurved leaf-trace than from an incurved one. The width of the back 
of the hook is the dictator of the width of the pinna-trace in the extramarginal 
supply ; the width of the marginal strip is the width of the whole margin of the 
unincurved leaf-trace. The back of a hooked leaf-trace is always "thin" as regards 
tracheides, and the pinna-traces which come off from it can give only a narrow strip 
of tracheides. Once the difficulty of carrying forward the water from pinna-departure 
to pinna-departure along the leaf was solved (by the development of hooks on the 
abaxial ends of the adaxial strands) the marginal type of supply must have been 
found an advantage rather than a drawback, because of its possibilities in supplying 
a long strip of tracheides to a departing pinna. This may explain its retention in 
the genus Polypodium, where the abaxial complications of the leaf-trace are often 
very elaborate and involved, and in some species of Pteris (P. cretica, P. umbrosa, 
etc.) where the abaxial arch and the lateral swellings of the leaf-trace are fairly 
prominent. The margins of the leaf-traces could be extended almost indefinitely to 



THE PINNA-TRACE IN THE FERNS. 375 

supply the pinnae, while the abaxial portion of the leaf-trace would cany forward the 
water-supply up the leaf. There is the possibility in the marginal type of " manu- 
facturing " so much more tracheidal tissue than is really necessary for the pinna- 
supply that no contraction would take place between the pinnae. But the actual 
number of tracheides does not seem to be altogether what keeps the water-supply in 
the proper direction. There is a deflection of the water-current towards the pinnae. 
It is really largely this deflection which has to be counterbalanced by the provision 
of an accessory set of tracheides which will carry the water forward. Where the 
pinnae are large the deflection will be great, and the counterbalancing hooks will be 
large ; where the pinnae are small the deflection is often negligible, and the hooks are 
small. This appears to lead to the conclusion that the Fern-leaf in phylogeny reduced 
its pinnae, the hooks of the leaf-trace disappearing in relation to this. When a 
large leaf appeared later, an accessory tracheidal system developed in order to 
counterbalance the deflections which would have drained the upper part of the leaf 
of its legitimate water-supply. 

All the features of the various leaf-traces cannot be fully explained on these 
theoretical grounds, but such distinctive characters of various leaf-traces as the 
incurved adaxial hooks connected with extramarginal supply to the pinnae, the 
reinforcement of the adaxial strand in Ferns with marginal supply, and the retention 
of the marginal type even by fairly large-leaved species can certainly be interpreted 
if we consider broadly the needs of the pinnae situated towards the tip of the leaf 
and the demand for water by large pinnae of leaves possessing leaf-traces with 
unincurved adaxial edges. 

A very interesting confirmation of the view that the adaxial or abaxial hooks of 
the adaxial portions of the leaf-trace serve to carry forward the water-supply past 
the pinna-departures is found in Salisbury's paper on " The Determining Factors in 
Petiolar Structure." He calls attention to Sinnott's conclusion {Annals, xxv. p. 187) 
that it is " extremely doubtful if the size of the transpiration current has had much 
influence on the development of the vascular supply," and confirms it for a species of 
Polypodium. There is a marked contraction of the xylem in a Fern-petiole, as 
Sinnott reiterates (pp. 187, 188), as we approach near to its point of junction with 
the stem. Salisbury, in emphasising the restriction of this contraction to a short 
distance at the base of the petiole, quotes (p. 266) the interesting observations of 
Ormsby, which show that the effect of contraction in water mains is negligible where 
the length of contraction is small as compared with the total distance traversed. 
But there is a possibility of contraction of the water-supply to a great part of the 
leaf at other points than just at the base. The departure of every pinna might 
cause such a contraction in the supply for the rest of the leaf. Only in a leaf of 
considerable length with a single large pinna (if we can imagine such a leaf) would 
it be possible to neglect the effect of the departure of the pinna-trace upon the 
supply for the rest of the leaf. The confirmation of Ormsby's rule is seen in the 



376 MR K. C. DAVIE ON 

elaborate precautions taken by leaf-traces to prevent undue contractions of their 
general water-supply due to the departures of successive pinnae. 

The two factors which have more than any others determined the development 
of the complications and simplifications of the margins of the Fern leaf-trace have 
been the need of carrying up past the earlier pinna-trace-departures a water-supply 
sufficient to meet the demands of the succeeding pinnae and the terminal portion of 
the leaf. The need for supplying the separate pinna-traces has been met by the 
giving-off of the parts of the leaf-trace nearest to those pinnae. As the leaves became 
larger and more heavily pinnate, provision was made for the water-supply of their 
increasing number by the development of the incurved hooks on the adaxial face of 
the leaf-trace. The departure of the pinna-traces took place still from the most 
conveniently situated part of the leaf-trace, but, as this was the back of a hook, the 
supply was extramarginal in origin. Then there seems to have been a condensation 
of the leaf- trace (perhaps in relation to a reduction in the size of the leaf), shown in 
the appearance of " broken " leaf-traces, either of the binary or of the many-stranded 
type. For some time the hooks were retained on the adaxial strands, but they 
gradually disappeared, and the pinna-supply was given off simply from the margins 
of the adaxial strands. Here and there arose again the need for carrying forward 
water past the pinna-trace departures. The marginal type of supply had become 
useful in its possibilities of extensive supply to the pinnae. And the difficulty of 
carrying forward the water-supply to the ultimate parts of the leaf was solved by 
pressing the abaxial strand or the abaxial portions of the adaxial strands into the 
service. Sometimes the adaxial strands were reinforced from the median strand as 
necessity arose ; at other times they were provided throughout their length with 
incurved abaxial hooks. If one may risk another generalisation from these details, 
it would seem that the Fern leaf in the course of its phylogeny had developed first 
in respect of length, at the same time as its appendages increased in size. Then there 
came a reduction both in the length of the leaf and in the size of the appendages, 
the reduction in length preceding the reduction in the size of the appendages. By 
that time the Fern leaf-trace had become thoroughly adapted to the needs of the 
leaf, and the later reductions or amplifications in special cases made but little change 
on the configuration of its adaxial portion, but mainly affected the abaxial part. 
Indeed, in the marginal type of pinna-supply which occurs in the more advanced 
Ferns we have the most improved type, and one which has proved to be the most 
adaptable. 

In conclusion, I desire to express my thanks to Professor I. Bayley Balfour, 
F.R.S., for granting me the privilege of obtaining most of the material 1 have 
examined from the Royal Botanic Garden, Edinburgh, and for communicating this 
paper; to Professor F. 0. Bower, F.R.S., and Professor D. T. Gwynne-Vaughan, 
F.L.S., for much helpful criticism ; and to Dr W. T. Gordon, F.R.S.E., for assistance 
in the preparation of the microphotographs. 



THE PINNA-TRACE IN THE FERNS. 377 



BIBLIOGRAPHY. 

Bertrand, C. Eg., et Cornaille, F. ('02), Etude sur quelques caracteristiques de la structure des Filicinees 

actuelles. 

• et P. Bertrand ('11), Le Tubicaulis Berthieri (sp. nov.), C. Eg. Bertrand et P. Bertrand. 

Bertrand, Paul ('09), Etudes sur lafronde des Zygopteridees, p. 127. 

('H)> Nonvelles remarques sur la fronde des Zygopteridees. 

Boodle, L. A. ('01), "On the Anatomy of the Gleicheniacese," Annals of Botany, xv. pp. 703-744. 

Bower, F. O. ('12), "Studies in the Phylogeny of the Filicales : II. LopJiosoria," Annals of Botany, xxvi. 

pp. 269-320. 
Chrysler, M. A. ('10), "The Nature of the Fertile Spike in the Ophioglossaceae," Annals of Botany, xxiv. 

pp. 1-18. 
Compton, R. H. ('09), " The Anatomy of Malonia sarmentosa, Baker," Neiv Phytologist, viii. pp. 299-309. 
Davie, R. O ('12), "The Structure and Affinities of Peranema and Diacalpe," Annals of Botany, xxvi. 

pp. 245-266. 

('13), "The Pinna-Trace in the Filicales," Report, British Association Meeting, 1913, p. 709. 

Gordon, W. T. ('11), "On the Structure and Affinities of Diplolabis Romeri (Solms)," Trans. Roy. Soc. 

Edin., xlvii., pt. iv., pp. 711-736. 
('11), "On the Structure and Affinities of Metaclepsydropsis duplex (Williamson)," Trans. Roy. Soc. 

Edin., xlviii., pt. i., pp. 163-190. 
Gwynne-Vaughan, D. T. ('01), "Observations on the Anatomy of Solenostelic Ferns : I. Loxsoma," Annals 

of Botany, xv. pp. 71-97. 

('03), " Observations on the Anatomy of Solenostelic Ferns, Part II.," Annals of Botany, xvii. pp. 689-740. 

('05), " On the Anatomy of Archangiopteris Henryi and other Marattiaceae," Annals of Botany, xix. 

pp. 259-271. 

■ ('11), "Some Remarks on the Anatomy of the Osmundaceae," Annals of Botany, xxv. p. 525. 

and Kidston, R. ('08). " On the Origin of the Adaxially-curved Leaf-trace in the Filicales," Proc. Roy. 

Soc. Edin., xxviii. pp. 433-436. 
"On the Fossil Osmundacese," Trans. Roy. Soc. Edin., Part I., vol. xlv. pp. 759-780; Part II., 

vol. xlvi. pp. 213-232 ; Part III., vol. xlvi. pp. 651-667 ; Part IV., vol. xlvii. pp. 455-476. 
Kidston, R. ('08), "On a New Species of Dineuron and of Botryopteris from Pettycur, Fife," Trans. Roy. 

Soc. Edin., vol. xlvi. pp. 361-364. 
Parmentier, P. ('99), "Recherches sur la structure de la feuille des Fougeres et sur leur classification," Ann. 

Sc. nat. Bot., 8 e ser., ix. pp. 289-361. 
Pelourde, F. ('06), "Recherches anatomiques sur la classification des Fougeres de France," Annates des 

Sciences nat. Bot., 9 e ser., iv. pp. 281-372. 
Salisbury, E. J. ('13), " The Determining Factors in Petiolar Structure," New Phytologist, xii. pp. 281-289. 
Seward, A. C. ('99), " On the Structure and Affinities of Matonia pectinata, R. Br.," Phil. Trans. Roy. 

Soc, B, vol. 191, pp. 171-209. 
Sinnott, E. W. ('10), "Foliar Gaps in the Osmundaceae," Annals of Botany, xxiv. pp. 113-115. 

('1 1), " The Evolution of the Filicinean Leaf-trace," Annals of Botany, xxv. p. 167. 

Tansley, A. G. ('08), Lectures on the Evolution of the Filicinean Vascular System. 

Thomae, K. ('86), "Die Blattstiele der Fame," Jahrb.f. wiss. Bot., xvii. pp. 136, 140 et seqq. 



TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 11). 52 



378 MR R. C. DAVIE ON THE PINNA-TRACE IN THE EERNS. 

EXPLANATION OF PLATES. 

All the figures are from untouched microphotographs. 

Plate XXXIII. figs. 1-8. 

Fig. 1. Transverse section of the petiole of Asplenium obtusatum, Forst., showing the departure of the 
vascular supply to the two basal pinnae, x 38. 

Fig. 2. Transverse section of the petiole of Davallia pallida, Mett., showing one pinna-trace nipped off 
from the leaf-trace, and the other in process of development. x 42. 

Fig. 3. Transverse section of the petiole of Aneimia hirta (L.) Sw., showing the departure of traces to the 
basal pinnae, x 40. 

Fig. 4. Transverse section of the petiole of Aneimia collina, Raddi. x 20. 

Fig. 5. Transverse section of the petiole of Loxsoma Gunninghami, R. Br., showing development of 
pinna-trace. x 30. 

Fig. 6. Transverse section of the petiole of Loxsoma Gunninghami, showing departure of pinna-trace, x 30. 

Fig. 7. Transverse section through one of the adaxial bundles of the petiole of Didymochl&na truncatula 
(Sw.) J. Sm., showing the development of the " extramarginal " type of vascular supply to the basal 
pinna. x 55. 

Fig. 8. Transverse section of the petiole of Leptopteris hymenophylloides, A. Rich., showing the departure 
of one of the pinna-traces. x 44. 

Plate XXXIV. figs. 9-15. 

Fig. 9. Transverse section of the petiole of Odontosoria chinensis (L.) J. Sm., var. Veitcliii, below the 
basal pinnae, showing the development of the pinna-traces. x 44. 

Fig. 10. Transverse section of the petiole of Hymenophyllum demissum (Forst.) Sw., showing one pinna- 
trace separated from the leaf-trace, and another in process of separation, x 42. 

Fig. 11. Transverse section of the petiole of Blechnum orientale, L., showing the development of one of 
the basal pinna-traces. x 40. 

Fig. 12. Transverse section through one of the strands of the leaf- trace of Gymnogramma Pearcei, 
Moore, var. robusta, showing the pinna-trace in process of being given off. x 40. 

Fig. 13. Transverse section of the petiole of Balantium eulcita (L'Herit.) Klf., showing the extension 
and buckling of the ends of the leaf-trace prior to the departure of the pinna-trace, x 20. 

Fig. 1 4. Transverse section of the petiole of Gleichenia flabellata, R. Br., showing the formation of the 
isolated cylindrical strands before the pinna-traces depart. x 33. 

Fig. 15. Transverse section of the petiole of Histiopteris incisa (Thbg.) J. Sm., showing the development 
of two pinna-traces. x 30. 

Plate XXXV. figs. 16-23. 

Fig. 16. Transverse section of the petiole of Histiopteris incisa (Thbg.) J. Sm., showing the completion 
of one of the pinna-traces. x 33. 

Fig. 17. Transverse section of the petiole of Botrychium lunaria (L.) Sw., showing the formation of one 
of the pinna-traces. x 33. 

Fig. 18. Transverse section of the petiole of Odontosoria retusa (Cav.) J. Sm., showing one pinna-trace 
free from the end of the leaf-trace, and the other in process of development. x 30. 

Fig. 19. Transverse section of the petiole of Onoclea sensibilis, L., showing the method of vascular supply 
to the basal pinnae. x 46. 

Fig. 20. Transverse section of one of the adaxial strands of the leaf-trace of Drynaria rigidula (Sw.) 
Bedd., showing the separation from it of a pinna-trace. x 53. 

Fig. 21. Transverse section of the petiole of GheilantJies myriophylla, Desv., var. elegans, showing the 
method of separation of the basal pinna-traces. x 33. 

Fig. 22. Transverse section of the petiole of Pteris cretica, L., showing the extension of the ends of the 
leaf-trace, prior to the departure of the basal pinna-traces. x 33. 

Fig. 23. Transverse section of the rachis of the leaf of Aneimia collina, Raddi, showing the method of 
vascular supply to the last two pinnae. x 42. 



is. Hoy. Soc. Edin r . 



Vol. L.— Plate XXXIII. 



Davie : Pinna — Trace in Ferns. 











fD. photo 



M'Farlane & Erskine. Eilin 



T: as. Roy. Soc. Edin r . 



Vol. L.— Plate XXXIV. 



Davie : Pinna— Trace in Ferns. 





10 






14 




15 



• photo. 



M'Farlane & Erekine, Edin. 



'Bins. Roy. Soc. Edin r 



Vol. L.— Plate XXXV. 



Davie : Pinna — Trace in Ferns. 




16 




17 





20 




21 




18 





19 



23 



3. photo 



M'Farlane & Erskine. Edin. 



( 379 ) 



XII. — Studies on the Pharmacological Action of Tetra-Alkyl-Ammonium 
Compounds. By Professor 0. R. Marshall. 

(Read December 15, 1913. MS. received March 24, 1914. Issued separately July 29, 1914.) 

II. THE ACTION OF TETRA-ETHYL-AMMONIUM CHLORIDE. 

The pharmacological action of tetra-ethyl-ammonium chloride has been investigated 
to a limited extent by Tillie,* Santesson and Koraen,! Tappeiner,} Boehm,§ and 
Hober and Waldenberg,|| in the course of other researches. Tillie, and Santesson 
and Koraen, describe its action on frogs ; Tappeiner merely states that it has 
no paralysing effect on the respiration of the rabbit ; Boehm, and Hober and 
Waldenberg, refer only to its effect on the isolated frog's muscle. The descriptions 
of the action of the iodide, which has been investigated by Jordan,! Brunton and 
Cash,* # Rothberger,! t Jacobj and Hagenberg,^ and Barger and Dale,§§ in the 
course of other work, are also somewhat meagre. Brunton and Cash, and Jacobj 
and Hagenberg, describe its action on frogs, and Brunton and Cash its action on 
rabbits ; but Jordan merely says that it produces on frogs a curare-like action but 
no muscarin effect, and Barger and Dale only state that it has no sympathomimetic 
action. The investigation of Rothberger was limited to its antagonistic action 
towards curarin. 

A point of some importance, which I have not seen mentioned, is the instability 
of the chloride when kept under ordinary conditions. It is not only hygroscopic, but 
it also decomposes and relatively quickly becomes acid. A specimen which had been 
kept in the laboratory in a well-stoppered bottle for over a year was found on 
analysis to contain 15 per cent, of water and 14'8 per cent, of hydrochloric acid. 
The remainder, as shown by a determination of the ethyl groups and the chloride 
content, was pure tetra-ethyl-ammonium chloride. Two other specimens obtained 
from reliable firms contained 4 '5 per cent, and 8 per cent, hydrochloric acid 
respectively when received. The commercial chloride, and this salt purified by 
dissolving in the smallest quantity of absolute alcohol, neutralising, filtering and 
drying, finally in a desiccator, were used in the investigation. 

General Effects. 

On Frogs. — The most prominent symptoms produced in grass frogs by injections 
into the dorsal lymph-sac were muscular contractions and trem'ors. Besides these, 

* Arch.f. exp. Path. u. Pharmak., xxvii. p. 17 [1890]. t Skand. Arch. f. Physiol, x. p. 225 [1900]. 

% Arch.f. exp. Path. u. Pharmak, xxxvii. p. 349 [1896]. § Ibid., lviii. p. 267 [1908]. 

|| Pftiiger's Arch., cxxvi. p. 337 [1909]. IT Arch.f. exp. Path. u. Pharmak, viii. p. 15 [1877]. 

** Trans. Roy. Soc. Lond., clxxv. p. 201 [1884]. tt Pftiiger's Arch., xcii. p. 424 [1902]. 

IX Arch.f. exp. Path. u. Pharmak, xlviii. p. 58 [1902]. " §§ Journ. of Physiol, xli. p. 28 [1910]. 

TRANS. ROY. SOC. EDIN., VOL. L. PART II. (NO. 12). 53 



380 PROFESSOR C. R. MARSHALL ON THE 

there were some loss of co-ordination, a varying degree of paralysis with diminution 
or absence of reflexes, and diminution or cessation of the respiratory movements. 
Winter frogs kept in the laboratory were less sensitive to this substance than 
recently caught summer frogs. 

The muscular contractions and tremors appeared in the case of summer frogs 
within three minutes of the injection of a large dose (2 mg. per gramme weight of 
frog), in five to eight minutes after a moderate dose (0*5 mg. per gramme weight of 
frog), and in twenty to thirty minutes after a minimal effective dose (0'05 mg. per 
gramme weight of frog). Most commonly they seemed to commence in the dorsal 
or abdominal muscles, but in some animals the muscles of the limbs were first 
influenced. Sometimes, and especially after large doses, the tremors and contractions 
were general from the outset. In all cases the irregularity in distribution and extent 
of the movements was marked. 

At first the contractions were usually fibrillary and shortly tonic in character. 
Later, large muscle-bundles or even whole muscles were involved, producing in the 
case of the limbs somewhat grotesque movements. Still later, if the dose administered 
were sufficient, they assumed the form of twitches, which became less frequent and 
finally ceased. In some cases they persisted for hours, and often continued after the 
cessation of all reflex activity. The movements of the hind-limb muscles were the 
last to disappear. Except after large doses, they were not constantly present during 
the earlier stages of the intoxication. Intervals of several minutes often occurred 
between successive twitches, and frequently they were induced by a voluntary 
movement. Such a movement generally intensified them if present. 

After doses larger than 0'3 mg. per gramme body-weight the stage of irregular 
muscular contractions usually passed into one of complete paralysis, which lasted a 
variable time. After the injection of 1 mg. per gramme of frog, paralysis occurred 
in fifteen minutes and complete recovery within eighteen hours. After double this 
dose, paralysis developed in five minutes and recovery did not occur. The muscles 
associated with respiration were the first to be paralysed and, if a non-lethal dose 
had been given, were the first to recover. After doses less than 0'3 mg. per gramme 
frog, complete paralysis did not follow, but more or less muscular weakness was 
noticeable. 

Tillie* states that doses of 0*01 g. to 0'02 g. induce in frogs powerful general 
fibrillary contractions before the onset of paralysis, but no mention is made of the 
occurrence of tremors by Santesson and KoRAEN.t They were, however, observed 
by Brunton and Cash \ and by Jacobj and Hagenberg § after the administration of 
the iodide. But Brunton and Cash || describe the symptoms produced by 0'007 g. 
to 0'028 g. of tetra-ethyl-ammonium iodide as being the same as those produced by 
similar doses of tetra-methyl-ammonium iodide. These were " spasmodic twitchings 
of trunk and limb muscles ; limbs drawn up in a very tremulous manner ; whole 

* Loc. cit. t Loc. cit. j Loc. cit., p. 205. t" Loc. cit., p. 58. || Loc. cit., p. 205. 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL- AMMONIUM COMPOUNDS. 381 

body twitches in response to pinching foot, even when the foot is no longer with- 
drawn ; death within a minute with larger dose." These symptoms, with the 
exception of the rapid death, have been present in all my experiments with effective 
doses of tetra-ethyl-ammonium chloride, but only on few occasions have I seen slight 
and transient fibrillary tremors in frogs after the injection of tetra-methyl-ammonium 
chloride. Tremors are, however, mentioned as occurring before the paralysis by 
Dufaux* and by Iodlbauer,! but they were not seen, although looked for, in the 
experiments of Santesson and Koraen,| and are not referred to by Tillie.§ In 
my experiments with tetra-methyl-ammonium chloride the symptoms produced by 
all doses from 0'006 mg. to 5 mg. per gramme weight of frog have been typically 
paralytic. And by injecting tetra-methyl-ammonium chloride previously or sub- 
sequently to the administration of tetra-ethyl-ammonium chloride I have prevented 
or annulled the contractions normally produced by the injection of the latter 
substance. Even if one-tenth the amount of the tetra-methyl salt be injected 
simultaneously with the tetra-ethyl compound, a notable limitation of the duration 
of the time the fibrillary contractions continue is seen. As will be shown later, the 
tremors produced by tetra-ethyl-ammonium chloride have their seat in the mechanism 
associated with the myo-neural junction, and it is possible that the different results 
obtained by different workers may have their explanation in the differences in this 
mechanism. To this point I hope to return in a later communication. I have, 
however, injected frogs at different times of the year and under different conditions 
with tetra-methyl-ammonium chloride without producing tremors. Nevertheless, 
fibrillary contractions may be produced for a brief period by applying small drops of 
a solution of tetra-methyl-ammonium chloride to an exposed muscle, and these 
commonly occur when an isolated muscle of a winter frog, but rarely when that of a 
summer frog, is immersed in a solution. 

In view of the statement of Brunton and Cash || that tetra-methyl-ammonium 
iodide and tetra-ethyl-ammonium iodide produce the same action, I have made a few 
experiments on frogs with these substances. If certain slight differences due to the 
iodide are ignored, the effects obtained are strictly comparable with those produced 
by the corresponding chlorides. Tetra-methyl-ammonium iodide proved much more 
powerful than tetra-ethyl-ammonium iodide, and it did not produce those powerful 
irregular contractions and tremors which characterise the action of the latter 
compound. Moreover, tetra-methyl-ammonium iodide, like the chloride, quickly 
caused diastolic arrest of the heart, at least in doses of 0'15 mg. per gramme of frog 
and above, an effect which was wanting in my experiments with tetra-ethyl- 
ammonium iodide. 

On Rabbits. — The only experiments dealing with the effects of the tetra-ethyl- 
ammonium compounds on mammals that I have been able to find are those of 

* Quoted by Santesson and Koraen, loc. cit. t Arch. Internat. de Pharmacod., vii. p. 188 [1900]. 

I Loc. cit, p. 220. § Loc. cit. || Loc. cit., p. 205. 



382 PROFESSOR C. R. MARSHALL ON THE 

Brunton and Cash * with tetra-ethyl-ammonium iodide, and those of Jacobj and 
Hagenberg t with tetra-ethyl-ammonium tri-iodide. Brunton and Cash, after 
injecting hypodermically one gramme of tetra-ethyl-ammonium iodide into a rabbit, 
observed trembling and shivering in eleven minutes, paresis of the fore part of the 
body in twenty minutes, cessation of the respiration with persistence of the corneal 
reflex in twenty-three minutes, and death in thirty-six minutes. After injecting 
0*5 g. into a rabbit, the animal lay with its head on the table twenty-one minutes 
after the administration, shaking from side to side. Five minutes later there were 
convulsive springing, lurching, and shuffling, with dyspnoea. Death occurred thirty 
minutes after the injection. Jacobj and Hagenberg state that hypodermic injections 
of O'l g. to 0*25 g. tetra-ethyl-ammonium tri-iodide per kg. body-weight suspended 
in glycerin, into cats, produced no symptoms. 

As the injection of 8 c.c. of a 2*5 per cent, solution of tetra-ethyl-ammonium 
chloride into the peritoneal cavity of a rabbit weighing 1975 g. produced no 
symptoms, intravenous medication was resorted to. The injections were made into 
one of the veins of the ear, and occupied about one minute. The lethal dose was 
found to be, for rabbits, about 0*05 g. per kg. body-weight. The intravenous 
injection of 0*02 g. per kg. merely caused slight dyspnoea and slight tremors of the 
fore part of the body a few minutes after the administration. Half this dose only 
caused slight deepening of the respiration. After the administration of a small 
lethal dose (0*06 g. per kg.) the animal remained normal and moved about for one 
minute ; then the respiration became slower and the heart-beats somewhat faster, 
and three minutes after the completion of the injection there appeared slight 
spasmodic tremors with head-nodding. Half a minute later a second more powerful 
tremor occurred, and twenty seconds later still more powerful tremors, which 
continued ten seconds. Further similar spasms were few and slight and transient. 
Five minutes after the termination of the injection the breathing was much slower 
and somewhat gasping in character, and the heart-beats appeared somewhat more 
forcible but were otherwise unchanged ; the animal sank on the table with the head 
to one side, and from time to time moved slowly. Ten minutes after the injection 
the respiration was still slower and air-hunger still more manifest ; the heart-beats 
were somewhat slower and weaker. The corneal reflex was present. From six to 
eight minutes later the animal made several spasmodic springs, after which the pupil 
dilated and the cornea became insensitive. The respiration ceased twenty and a half 
minutes after the cessation of the injection, and the heart-beats a few seconds later. 

The injection of 0*1 g. per kg. intravenously produced deep convulsive breathing 
before the injection was completed. Erection of the tail and convulsive movements 
of the hind limbs, which lasted for a few seconds, quickly followed, and the animal 
then lay on its side paralysed. The corneal reflex was absent, but a few gasping 
movements of the mouth occurred, the last about two minutes after the termination 

* Loc. cit., p. 210. t Loc. cit., p. 60. 



PHARMACOLOGICAL ACTION OF TETRA-ALKYL- AMMONIUM COMPOUNDS. 383 

of the injection. The heart-beats were apparently normal at this time, but later 
they became weaker and less frequent, and finally ceased seven minutes after the 
end of the injection. Two minutes previously a few tremors of the right fore limb 
were noticed. 

After the intravenous injection of 0'25 g. per kg. body- weight, death occurred 
within a minute of the termination of the injection. A few slight convulsive move- 
ments, chiefly of the hind limbs, occurred during the injection, but the breathing 
had ceased and the fore part of the body was completely, and the hinder part almost 
completely, paralysed at the end of the injection. The heart-beats were then feeble ; 
they ceased before another minute had passed. The pupils were markedly contracted. 
Tremors of the back muscles occurred soon after the cessation of the injection, and 
somewhat later tremors of the abdominal muscles. Half an hour after death tremors 
in the hind limbs were present. 

Effect on Nerves and Muscles. 

Both the muscular contractions and the paralysis observed in frogs are peripheral 
in origin. They are obtained as easily in frogs in which the brain and spinal cord 
have been pithed as in normal animals, and are readily obtained by the local applica- 
tion of the substance to exposed muscles. 

Santesson and Koraen # state that they found it difficult to determine the 
paralysing action of tetra-ethyl-ammonium chloride on the nerve-endings owing to 
its action on the muscle-substance. This difficulty has not presented itself in my 
experiments. In all, whether on intact frogs or on muscle-nerve preparations, the 
myo-neural junction was found to be paralysed at a time when the irritability of the 
muscle was little affected. In a frog which had received 1*5 mg. per gramme body- 
weight into the dorsal lymph-sac, and which, after the development of preliminary 
tremors, was completely paralysed in ten minutes, the muscles when tested twenty 
hours later were found to react with the secondary coil (the primary current being 
obtained from one accumulator cell) at 19 cm., while the sciatic nerves did not 
react with the coil full up. In a second experiment, in which the frog received 2 mg. 
per gramme body-weight and was pithed one and three-quarter hours after the 
injection, the gastrocnemii muscles reacted with the secondary coil at 40 cm., the 
sciatic nerves being unirritable with the strongest current. With smaller doses the 
same phenomena but in less degree were observed. After the injection of 0'4 mg. 
per gramme of frog and pithing three hours later, the gastrocnemii reacted with the 
secondary coil at 36 cm., the right sciatic was irritable with the coil at 8 "5 cm., and 
the left sciatic with the coil at 4'5 cm. 

By means of a Claude Bernard experiment the predominant influence of tetra- 
ethyl-ammonium chloride on the nerve-endings is more decidedly shown. Thus, in a 
frog with the brain pithed and with the left iliac artery and vein ligatured and 

* hoc. cit., p. 225. 



384 PROFESSOR C. R. MARSHALL ON THE 

two ligatures round the upper part of the left thigh, the injection of 0"8 mg. per 
gramme of frog produced the following result when the nerves and muscles were 
tested nineteen hours later. The numbers refer to the position of the secondary 
coil : — 

Left gastrocnemius muscle, 18 '5 cm. Left sciatic nerve, 17 cm. 

Right gastrocnemius muscle, 17 cm. Right sciatic nerve, not irritable. 

In another experiment, in which the right thigh was ligatured, the muscles and 
nerves were tested two and a half hours after the injection, with the following 
result : — 

Right gastrocnemius muscle, 20"5 cm. Right sciatic nerve, 22'5 cm. 

Left gastrocnemius muscle, 19 cm. Left sciatic nerve, not irritable. 

If a dose insufficient to produce paralysis is given, the nerves may be found as 
irritable as the muscles, or may be found as irritable as the muscles on one side and 
less irritable on the other. Thus, in a frog to which O'l mg. per gramme body-weight 
was given, the following results were obtained when the animal was pithed two 
hours after the administration : — 

Left gastrocnemius muscle, 44'5 cm. Left sciatic nerve, 43*5 cm. 

Right gastrocnemius muscle, 44*5 cm. Right sciatic nerve 20 cm. 

But in these cases it was found that continued stimulation of the nerve induced 
exhaustion more quickly than normally. The difference on the two sides, in those 
cases in which it was present, was in all probability due to greater diffusion of 
some of the solution along the lymph -sacs of one side, and consequently a greater 
local effect. 

The greater paralysing influence on the nerve-endings as compared with the 
muscle may also be shown by means of a muscle-nerve preparation. When such a 
preparation is steeped in an isotonic solution of tetra-ethyl-ammonium chloride, the 
nerve is found to be unirritable before the irritability of the muscle is much affected. 
And as the application of an isotonic solution to the nerve-trunk does not materially 
influence the irritability of the nerve, the paralysing action of the substance must be 
attributed to an action on the so-called nerve-endings. 

The irregular mu